M. Gargaud & Al. 2015 - Encyclopedia of Astrobiology PDF

Muriel Gargaud
William M. Irvine
Editors-in-Chief Daniel Rouan
Ricardo Amils Tilman Spohn
José Cernicharo Quintanilla Stéphane Tirard
Henderson James Cleaves II Michel Viso
Daniele L. Pinti Editors
Encyclopedia of
Astrobiology
Second Edition
1 3Reference
Encyclopedia of Astrobiology
Muriel Gargaud • William M. Irvine
Editors-in-Chief
Ricardo Amils • Henderson James (Jim)

Cleaves II • Daniele L. Pinti
José Cernicharo Quintanilla
Daniel Rouan • Tilman Spohn
Stéphane Tirard • Michel Viso
Editors
Encyclopedia of
Astrobiology
Second Edition
With 703 Figures and 99 Tables

Editors-in-Chief
Muriel Gargaud William M. Irvine
CNRS-Universite de Bordeaux University of Massachusetts
Laboratoire d’Astrophysique Amherst, MA, USA
de Bordeaux
Floirac, France
Editors
Ricardo Amils Henderson James (Jim) Cleaves II
Departamento de Biologia Molecular Earth–Life Science Institute (ELSI)
Universidad Autónoma de Madrid Tokyo Institute of Technology
Madrid, Spain Meguro–ku, Tokyo, Japan
Daniele L. Pinti José Cernicharo Quintanilla

GEOTOP Research Center for Department of Astrophysics
Geochemistry and Geodynamics Laboratory of Molecular Astrophysics
Université du Québec à Montréal Iorrejón de Ardoz
Montréal, QC, Canada Madrid, Spain
Daniel Rouan Tilman Spohn

LESIA, Observatoire Paris-Site Deutsches Zentrum f€
ur Luft- und
de Meudon Raumfahrt (DLR), Institut f€
ur
Meudon, France Planetenforschung
Berlin, Germany
Stéphane Tirard
Centre François Viéte d’Histoire Michel Viso
des Sciences et des Techniques EA 1161 CNES/DSP/SME, Vétérinaire/DVM
Faculté des Sciences et des Astro/Exobiology
Techniques de Nantes Paris Cedex 1, France
Nantes, France
ISBN 978-3-662-44184-8 ISBN 978-3-662-44185-5 (eBook)

ISBN 978-3-662-44186-2 (print and electronic bundle)
DOI 10.1007/978-3-662-44185-5
Library of Congress Control Number: 2015947250
Springer Heidelberg New York Dordrecht London

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Foreword to the First Edition
Are we alone? Long an object of speculation or fiction, if not heresy, this

question entered the field of science on November 1, 1961, at the National
Radio Astronomy Observatory in Green Bank, Virginia, where a number of
scientists, including Melvin Calvin, who had just been awarded the Nobel
prize in chemistry for his work on photosynthesis, and the charismatic Carl
Sagan, gathered at the invitation of a young astronomer, Frank Drake, to
launch the Search for Extra-Terrestrial Intelligence (SETI) project. Since
then, batteries of increasingly powerful radiotelescopes have been scanning
space for messages sent out by some extraterrestrial civilization, so far
in vain.
At the same time, in the wake of widening space exploration, a new
discipline was born that has the distinctive peculiarity of having three
names – exobiology, astrobiology, and bioastronomy – and no as-yet-known
object. The purpose of this new discipline is more modest than that of the
SETI project: to detect signs of extraterrestrial life, not necessarily intelligent.
To guide this quest, we have available vast knowledge that has been gained
in the last few decades concerning the basic mechanisms of life. This knowl-
edge, in turn, has illuminated our concept of the origin of life. Even though we
do not know how or under what conditions this phenomenon took place, we
may safely affirm that if life arose naturally, which is the only scientifically
acceptable assumption, its origin must have depended on “chemistry.” By its
very nature, chemistry deals with highly deterministic, reproducible events
that are bound to take place under prevailing physical–chemical conditions. If
even a very slight element of chance affected chemical reactions, there would
be no chemical laboratories, no chemical factories. We could not afford
the risk.
A conclusion that emerges from this consideration is that life, as a product
of environmentally enforced chemistry, was bound to arise under the
physical–chemical conditions that prevailed at the site of its birth.
This statement, at least, holds true for the early steps in the origin of life,
until the appearance of the first replicable substance, most likely RNA. Once
this happened, “selection” became added to chemistry, introducing an ele-
ment of chance in the development of life. Contrary to what has often been
claimed in the past, this fact does not necessarily imply that the process was
ruled by contingency. There are reasons to believe that, in many instances,
v
vi Foreword to the First Edition
chance provided enough opportunities for selection to be optimizing and,

therefore, likewise obligatory under prevailing conditions.
Thus, in so far as chemistry and optimizing selection played a dominant
role in the process, the development of life appears as the obligatory outcome
of prevailing conditions. Hence the assumption that the probability of the
appearance elsewhere in the universe of forms of life resembling Earth life in
their basic properties is approximately equal to the probability of the occur-
rence elsewhere in the universe of the physical conditions obtained at the site
where Earth life arose.
In the eyes of many astronomers, this probability is very high. It is
estimated that some 30 billion sun-like stars exist in our galaxy alone and
that the total number of galaxies in the universe is on the order of 100 billion.
This means, to the extent that our galaxy may be taken as a representative
sample of galaxies in general, there may be some 3,000 billion sun-like stars
in the universe. Unless our solar system should be the product of extremely
unlikely events, the probability of there being planets similar to Earth (or to
whatever celestial object served as the cradle of Earth life) seems very strong.
Recent findings are most encouraging in this respect, by revealing that
planet formation is not a rare event, with more than 400 planets already
identified around a number of nearby stars. Although no habitable Earthlike
extrasolar planet has yet been found, this may be partly due to technical
limitations. The prospects that, with improved technologies, such a planet
may be discovered some time in the future are far from negligible. Signs of
life on such a planet, although more difficult to detect, may likewise yield
technological progress.
As by now, the enormous research effort expended within the framework
of the new discipline of exobiology-cum-bioastronomy-cum-astrobiology
has already produced a wealth of new findings, in fields ranging from physics
and cosmology to chemistry, biochemistry, and molecular biology. These
findings have provided rich material for this Encyclopedia. Its editors and
authors are to be commended for making this material widely available in
easily accessible form.
14 January 2011 Christian de Duve
References
1. de Duve C (2005) Singularities landmarks on the pathways of life. Cambridge Univer-
sity Press, New York
Preface to the First Edition
Where do we come from? Are we alone in the Universe? Where are we going?
These are the questions addressed by astrobiology – the study of the origin,
evolution, distribution, and the future of life in the Universe.
Encyclopedias are unusual works. A quote from the prologue of one of the
more famous early encyclopedias is instructive:
“. . .the purpose of an encyclopedia is to collect knowledge disseminated
around the globe; to set forth its general system to the men with whom we
live, and transmit it to those who will come after us, so that the work of
preceding centuries will not become useless to the centuries to come; and so
that our offspring, becoming better instructed, will at the same time become
more virtuous and happy, and that we should not die without having rendered
a service to the human race in the future years to come”. Diderot and
d’Alembert, Encyclopédie (1751).1
Diderot and d’Alembert’s eighteenth century Encyclopédie was indeed
ground-breaking, but perhaps more remarkable is the degree to which their
description resembles the modern concept of genetic inheritance and natural
selection: a civilization’s accumulated knowledge being analogous to the
traits encoded in an organism’s time-tested DNA genome. In many ways,
the Encyclopédie addressed the goals of astrobiology; between the lines, we
find aspects of what makes biology biology.
Encyclopedias have now existed for approximately 2,000 years, the first
being Pliny the Elder’s Naturalis Historia, which was a compendium of the
knowledge available to a citizen of the Roman Empire as documented by the
first century AD.2 It contained ~ 20,000 facts from 2,000 sources written by
200 authors. The present volume contains an unknown number of “facts”
(indeed, some of the content will likely be proven false, as science is a living,
breathing accumulation of presently accepted knowledge, all subject to future
revision), but it does include more than 1,700 contributions, references
uncounted thousands of prior publications, and is written by 385 authors.
Modern encyclopedias are derived from the dictionaries of the eighteenth
century. The two are similar in that both are arranged alphabetically and
generally are the work of a team of expert contributors. They differ in that
encyclopedias contain a deeper level of analysis of the included terms and
attempt to cross-reference and place the assembled contents in a useful context.
The first encyclopedias attempted to cover all human knowledge. This is
now impossible for a printed work because the body of human knowledge is
vii
viii Preface to the First Edition
presently growing exponentially, with no end in sight. Encyclopedias now

exist for almost every definable field of study. A field requires a certain degree
of maturity to have an encyclopedia, and conversely, the publication of an
encyclopedia commonly records the birth of a definable field of study.
Astrobiology is an interdisciplinary field, spanning geology, chemistry, phys-
ics, astronomy, biology, engineering, and computer science, to name only the
core fields of study.
While some of these fields of research are fairly well mapped, many others
are in rapid flux, and still others remain perennially enigmatic, awaiting future
breakthroughs by the scientists of tomorrow. To this end, the Encyclopedia of
Astrobiology is primarily aimed at younger scientists or scientists new to the
field who wish to understand how their expertise coincides with current
knowledge in other areas of study. It is hoped that the encyclopedia will
serve to orient researchers to the current state of the art. A more in-depth
discussion of many of the topic areas can be obtained by referring to college
or graduate level texts or to the articles cited at the end of many of the entries.
Encyclopedias are snapshots of the state of knowledge at a particular time.
In 1844, the book Vestiges of the Natural History of Creation was published
anonymously (it was later found to have been written by Scottish publisher
William Chambers) and created a public sensation.3 It offered a sweeping and
very secular view of the development of the Solar System, stretching from the
nebular hypothesis to the development of man. While primitive by modern
standards (it was, after all, based on state-of-the-art early nineteenth century
science), it was in many ways remarkably similar to modern cosmology. In
broad brushstrokes, it is the precursor to the worldview developed in Carl
Sagan’s Cosmos4 and the grand view of myriads of habitable planets implicit
in the Drake equation. The implication of Vestiges was simply this: the
Universe operates everywhere and at all times according to physical princi-
ples, and the evolution of matter is largely predictable and often progressive,
proceeding from the simple to the complex.
Science has advanced dramatically since Chambers’ book was published.
It is truly a long way from Sir William Hershel’s 40-ft telescope to the
Herschel Space Telescope,5 and from a Universe with seven known planets
orbiting the Sun to one with more than 500 planets orbiting other stars. It is
also a long way from the work of Black, Priestly, and Lavoisier6 to SELEX
technology and high-throughput automated chemical screening and analysis,
and from Lyell’s Principles of Geology7 to plate tectonics and isotope geo-
chemistry. Nonetheless, certain questions permeate the sciences across time
and discipline. Woese’s three domains of life8 are direct descendents of
Linnaeus’ early classification scheme, and both are attempts to unify and
classify terrestrial organisms. Darwinism has offered an underlying mecha-
nism for doing so that has allowed for unification of the assorted observations
of the living world. However, the question of whether terrestrial life is unique
in the universe has fascinated mankind for millennia.
It was not until 1959, when NASA began funding the search for life in the
Universe in its Exobiology program, that we at last achieved the technological
prowess to try to answer this question.9 The paleontologist George Gaylord
Preface to the First Edition ix
Simpson famously noted shortly thereafter that Exobiology was a science

“that has yet to demonstrate that its subject matter exists.”
NASA’s first exobiology grant was awarded to Wolf Vishniac for the
construction of the Wolf Trap, a device for detecting bacteria on Mars. Due
to size limitations, the device never flew, but various descendants have made
the trip to Mars and returned various negative or tantalizingly ambiguous
results. These results are, amusingly, either disappointingly or encouragingly
ambiguous, depending on one’s point of view. Despite remarkable progress in
the sciences, humanity still has no answer to the question, “Are we alone?,”
though the question is in principle answerable. The search continues
enthusiastically.
Why should we think there might be life elsewhere in the Universe? In
1960, the radio-astronomer Frank Drake developed his now-famous equation
for estimating the number of communicating civilizations in the Galaxy:
N ¼ R f p ne f e f i f c L;
where N is the number of civilizations in our galaxy for which communica-

tion might be possible, R is the average rate of star formation per year, fp is
the fraction of stars that have planets, ne is the average number of planets that
can support life per star with planets, f‘ is the fraction of the planets that can
support life on which life actually develops, fi is the fraction of those on which
intelligent life develops, fc is the fraction of those on which civilizations
communicate using detectable signals, and L is the length of time these
civilizations communicate.
When Drake unveiled his equation in 1960 and estimated that there were
maybe ten communicating civilizations in the Galaxy, few of the parameters
were known with any certainty; the rate of star formation was perhaps the
only solid measurable value. Fifty years later, the flourishing search for
exoplanets has placed the focus on the second value (notably, it now appears
to be close to what Drake estimated, ~50 %). Hundreds of exoplanets have
been found around other stars, and current technology allows the observation
of even small planets. Theory suggests that the fraction of stars with Earthlike
planets is somewhere near 10 % (again, surprisingly, and a tribute to back-of-
the-envelope calculations, not far from Drake’s initial estimate).
The least well-known value is the question of how difficult is it for life to
begin (one of the “perennially enigmatic” facts mentioned above). Based on
present knowledge, the fraction of planets on which life actually emerges
(fl) could be anywhere from very, very close to 0 or far closer to 1. We simply
do not know. On the ends of the spectrum, the scientific community is divided
into two equally “hunch-”based camps: first, life is inevitable and is a cosmic
imperative (where conditions are appropriate) and, second, the origin of life
requires such a concatenation of improbable events that it is the scientific
equivalent of a miracle.
On the one planet we know of with life, our own, putative evidence in the
form of isotopically light carbon appears in the earliest known sedimentary
x Preface to the First Edition
rocks, suggesting life emerged relatively early in the history of the planet,
although we do not know whether this took place 100 years or 700 million
years after the planet formed. This implies that either something extraordi-
nary happened on Earth, or that the origin of life is a mundane phenomenon
on young planets, given appropriate chemistry, environmental conditions,
and enough time. Radioastronomy has provided a glimpse of the chemical
inventory of the cosmos which does appear to be universal. Spectral signa-
tures of a veritable zoo of organic compounds suggest that the Universe is
strewn with the potential precursors of life. Organic carbon (in the form of
carbon monoxide) has now been observed as far back as 13 billion years ago,
only some 700 million years after the birth of the Universe in the Big Bang.
The picture emerging, reminiscent of Chambers’ universe, is that physics and
chemistry are the same everywhere in the Universe, and that the Earth,
although remarkable in many respects, may not be unique.
As in any factorial equation, the most important values are the ones with
the largest uncertainty. Two approaches could shed light on the “fl problem”:
the duplication of the process in the laboratory or the discovery of life on
another planet. It is difficult to say whether the first approach will ever
succeed to anyone’s complete satisfaction, given that the origin of life on
Earth was a historical event that happened when no one was around to witness
it. The second approach, while fraught with technological difficulties, is
perhaps more promising. To that end, numerous instruments and space mis-
sions have been designed and launched to explore the Solar System and
beyond. The spectral signatures of planets around nearby stars are being
monitored for the characteristic signs of life such as the signature of disequi-
librium chemistry in the form of the presence in their atmospheres of both
oxidized and reduced gases.
While the answers to the vast questions that define astrobiology as a field
of study are unclear, it is evident that answering them will require an
interdisciplinary effort, stretching across international borders. One is hesi-
tant to speculate what the answer to the question, “Are we alone?” will
ultimately be. As good scientists, we should probably withhold judgment
until the data are in. As better scientists, we must join hands and find the data.
The editors of the Encyclopedia of Astrobiology hope that this volume will
contribute to this effort.
The Editors
Notes
1. A complete English and French version of the Encyclopédie can be found at http://quod.
lib.umich.edu/d/did/
2. For a complete English translation of Pliny the Elder’s The Natural History by John
Bostock see http://www.perseus.tufts.edu/hopper/text?doc=Plin.+Nat.+toc&redirect=
true. A complete Latin version can be found at http://www.perseus.tufts.edu/hopper/
text?doc=Perseus:text:1999.02.0138:toc&redirect=true
3. Chambers R (1994) Vestiges of the natural history of creation and other evolutionary
writings. University of Chicago Press
Preface to the First Edition xi
4. Cosmos was a remarkable 13-part popular science series narrated by Carl Sagan which
aired in 1980. Most if not all of the episodes can be viewed on line, and a book was spun
off: Sagan C (1985) Cosmos. Ballantine Books
5. For a survey of the early developments in astronomy, see Lankford J (ed) (1996) History
of astronomy: an encyclopedia, 1st edn. Routledge
6. For an excellent discussion of the early history of chemistry (including the work of
Black, Priestly and Lavoisier) see Partington JR (1989) A short history of chemistry, 3rd
revised edn. Dover Publications
7. Lyell C (2010) Principles of geology: being an inquiry how far the former changes of the
earth’s surface are referable to causes now in operation. Nabu Press (March 1, 2010).
Originally published in three volumes between 1830–1833
8. Woese C, Kandler O, Wheelis M (1990) Towards a natural system of organisms:
proposal for the domains Archaea, Bacteria, and Eucarya. Proc Nat Acad Sci USA
87(12): 4576–4579
9. For an insightful recounting of the early history of NASA’s early efforts in exo- and
astrobiology (including discussion of the roles of Wolf Vishniac and Frank Drake) see
Dick SJ, Strick JE (2005) The living universe: NASA and the development of astrobi-
ology. Rutgers University Press
Preface to the Second Edition
The Encyclopedia of Astrobiology serves as the key to a common under-

standing of the astrobiology field among astronomers and astrophysicists,
biologists and biochemists, chemists, geologists and geochemists, space
scientists, historians of science, and others working in this interdisciplinary
and rapidly expanding field. In the past few years, the pace of advancement in
astrobiology has become so rapid that we have felt it necessary to update and
expand many of the earlier entries and to augment the original edition of this
Encyclopedia, published in 2011, with some 300 new entries. Exciting results
include the discovery of some 1,200 planets (as of early 2015) orbiting stars
other than the Sun, some with masses close to that of Earth; the continuing
exploration of the conditions possibly relevant to life on Solar System bodies
such as Mars, Saturn’s satellites Titan and Enceladus, and comets; the in vitro
evolution of increasingly capable and diverse ribozymes; the progress toward
synthetic cellular systems that can undergo Darwinian evolution; the discov-
ery of increasingly complex organic molecules in interstellar space; and the
recently launched and newly planned space missions and telescopes that will
further our knowledge of the universe and make significant contributions to
the goals of astrobiology. All of this research is relevant to our increasingly
detailed understanding concerning the origin of life on our own planet and the
possible occurrence of life elsewhere in the universe. Finally, the History of
Science section is expanded to include speculation on topics such as the origin
of species and the possibility of inhabited planets beyond the Earth by
classical and medieval scholars, the latter from both Europe and the Islamic
world.
We believe that both new and experienced researchers as well as graduate
students – either in the adjacent fields of astrobiology or those new to the
subject – will appreciate this reference work during their quest to understand
the whole picture. To aid this process, we introduce in this edition a Table of
Contents, broken down by research area, which nicely illustrates the breadth
and the depth of the field of astrobiology. Although members of the different
disciplines commonly employ their own terminology and technical language,
here we have made a special effort to eliminate specialized jargon and
overtechnical terms from the Encyclopedia. Synonyms and keywords from
xiii
xiv Preface to the Second Edition
the 1st edition have been carefully revisited, with many newly added and
redundant terms deleted.
Both the carefully selected group of active researchers contributing to this
work as well as the expert field editors hope that this new edition will be
valuable to the scientific community and accelerate the interdisciplinary
advance of astrobiology.
The Editors
Acknowledgement to the Second Edition
A brief note is warranted about how we constructed the Encyclopedia.

A glossary of terms was first compiled by a team of experts in each field. It
was then cross-referenced between fields to check for conceptual overlap and
was then both expanded and pared down to produce a consensus entry list.
Authors with peer-recognized contributions to their fields of study were
then invited to contribute entries appropriate to their expertise. After a final
draft was submitted, entries were proofread and vetted for scientific accuracy
and readability by a team of field editors, then edited and modified to be
accessible by a reader with general knowledge of college-level science.
Finally, the entries were cross-referenced and edited for stylistic consistency
and ease of reading.
The editors would like to sincerely thank all the authors of the content of
the Encyclopedia for their efforts and understanding throughout the long and
at times difficult triple review process. We are particularly grateful to those
who also accepted to act as nonspecialist reviewers for fields other than
their own.
We would also like to thank several people who, although not authors,
served as external reviewers for a significant number of entries: Maxence
Claeys (Ecole Centrale Paris, France), Carlos Garcia-Ferris (Universitat de
València, Spain), David Hochberg (CAB, Madrid, Spain), Pierre Léna
(Académie des Sciences, Paris, France), Susan Leschine (University of Mas-
sachusetts Amherst, USA), and Jean Vandenhaute (University of Namur,
Belgium).
We express our gratitude to our respective institutions, especially those
who facilitated and aided in the organization and funding of editorial meet-
ings: Centre National de la Recherche Scientifique (CNRS, France), Centre
National d’Études Spatiales (CNES, France), Laboratoire d’Astrophysique de
Bordeaux (France), Université Bordeaux 1 (France), GEOTOP Research
Center for Geochemistry and Geodynamics (Université de Québec à Mon-
tréal, Canada), Natural Sciences and Engineering Research Council of Can-
ada, European Science Foundation (Archean Environment Research
Networking Program), Centro de Astrobiologı́a (CAB, INTA-CSIC, Madrid,
Spain), Universidad Autónoma de Madrid, NASA Goddard Space Flight
Center Cooperative Agreement NNX09AH33A with the University of Mas-
sachusetts. This work was also supported by the European COST Action TD
xv
xvi Acknowledgement to the Second Edition
1308 “Origins and evolution of life on Earth and in the Universe” and the
Erasmus + European Astrobiology Campus.
Last, but certainly not least, we express our sincere appreciation to the
editorial staff of Springer, in particular Saskia Ellis and Daniela Graf, who
lent technical and administrative support throughout the entire process.
The Editors
Editors-in-Chief
Muriel Gargaud CNRS-Universite de Bordeaux, Laboratoire

d’Astrophysique de Bordeaux, Floirac, France
William M. Irvine University of Massachusetts, Amherst, MA, USA
xvii
Field Editors
Ricardo Amils Departamento de Biologia Molecular, Universidad

Autónoma de Madrid, Madrid, Spain
Field: Life Sciences
Henderson James (Jim) Cleaves II Earth–Life Science Institute (ELSI),
Tokyo Institute of Technology, Meguro–ku, Tokyo, Japan
Field: Chemical Sciences
Daniele L. Pinti GEOTOP Research Center for Geochemistry and
Geodynamics, Université du Québec à Montréal, Montréal, QC, Canada
Field: Earth Sciences
José Cernicharo Quintanilla Department of Astrophysics, Laboratory of
Molecular Astrophysics, Iorrejón de Ardoz, Madrid, Spain
Field: Astrophysics and Astrochemistry
Daniel Rouan LESIA, Observatoire Paris-Site de Meudon, Meudon, France
Field: Astrophysics and Astrochemistry
Tilman Spohn Deutsches Zentrum f€ur Luft- und Raumfahrt (DLR), Institut
f€ur Planetenforschung, Berlin, Germany
Field: Planetary Sciences and Exoplanets
Stéphane Tirard Centre François Viéte d’Histoire des Sciences et des
Techniques EA 1161, Faculté des Sciences et des Techniques de Nantes,
Nantes, France
Field: History and Philosophy of Science
Michel Viso CNES/DSP/SME, Vétérinaire/DVM, Astro/Exobiology, Paris
Cedex 1, France
Field: Space Missions, Ground Facilities and Planetary Protection
and
Institutions and Organizations
xix
Section Editors
Alessandro Airo Institut f€ur Geologische Wissenschaften Tektonik und

Sedimentäre Geologie, Freie Universität Berlin, Fachbereich
Geowissenschaften, Berlin, Germany
Section: Inner Solar System
Francis Albarède Ecole Normale Supérieure de Lyon, Lyon, France

Section: Early Earth Geochemistry
Yann Alibert Space Research and Planetary Sciences, Physics Institute,

University of Bern, Bern, Swiss
Section: Planetary Formation and Dynamics
Philippe André Laboratoire AIM, IRFU/Service d’Astrophysique, CEA

Saclay, Gif-sur-Yvette, France
Section: Stars – Formation and Evolution
Kristin Bartik Université libre de Bruxelles, Brussels, Belgium

Section: Chemistry – General Definitions
Hugues Bersini IRIDIA, Université Libre de Bruxelles, Brussels, Belgium

Section: Artificial Life
Carlos Briones Centro de Astrobiologı́a (CSIC/INTA), Consejo Superior de

Investigaciones Cientı́ficas, Torrejon de Ardoz, Madrid, Spain
Section: Genetics and Evolution
Therese Encrenaz LESIA, Observatoire de Paris - Section de Meudon,

France
Section: Outer Solar System
Fernando B. Figueiredo CITEUC, University of Coimbra, Coimbra,

Portugal
Section: History and Philosophy of Science
xxi
xxii Section Editors
Felipe Gomez Centro de Astrobiologı́a (CSIC/INTA), Instituto Nacional de

Técnica Aeroespacial, Torrejón de Ardoz, Madrid, Spain
Section: Biology – General Definitions
John Lee Grenfell German Aerospace Center (DLR), Berlin, Germany
Section: Planetary and Exoplanetary Atmospheres
Nader Haghighipour Institute for Astronomy, University of Hawaii-
Manoa, Honolulu, Hawaii, HI, USA
Section: Exoplanetary Systems
Ravit Helled Geophysical, Atmospheric and Planetary Sciences, Tel Aviv
University, Raymond and Beverly Sackler Faculty of Exact Sciences,
Tel Aviv, Israel
Section: Planetary Formation and Dynamics
Christoph Heubeck Institut f€ur Geowissenschaften, Friedrich-Schiller-
Universität Jena, Jena, Germany
Section: Archean Geology
Gerda Horneck DLR German Aerospace Center, Institute of Aerospace
Medicine, Radiation Biology, Köln, Germany
Section: Microbiology in Space
Emmanuelle J. Javaux Palaeobiogeology-Palaeobotany-Palaeopalynology,
Geology Department, Université de Liège, Liège, Belgium
Section: Traces of Life
Kensei Kobayashi Yokohama National University, Tokiwadai, Hodogaya-ku,
Yokohama, Japan
Section: Chemistry – General Definitions
Juli Peretó Institut Cavanilles de Biodiversitat i Biologia Evolutiva,
Universitat de València, València, Spain
Section: Biochemistry
Andrew Pohorille NASA Ames Research Center, Mountain View, CA, USA
Section: Prebiotic Chemistry and Origins of Life
Nikos Prantzos Institut d’Astrophysique de Paris, Paris, France
Section: Nucleosynthesis
Barbara Stracke Deutsches Zentrum f€ur Luft- und Raumfahrt (DLR),
Institut f€
ur Planetenforschung, Berlin, Germany
Section: Inner Solar System
Contributors
José Pascual Abad Facultad de Ciencias, Departamento de Biologı́a Molec-

ular, Universidad Autónoma de Madrid, Cantoblanco, Madrid, Spain
Farah Abdul-Rahman University of Massachusetts Amherst, Amherst,
MA, USA
Delphine Acolat Centre François Viète, Université de Bretagne
Occidentale, Quimper, France
Angeles Aguilera Laboratorio de Extremófilos, Centro de Astrobiologı́a
(INTA-CSIC), Torrejón de Ardoz, Madrid, Spain
Marcelino Agundez Instituto de Ciencia de Materiales de Madrid, Madrid,
Spain
Alessandro Airo Institut f€ur Geologische Wissenschaften Tektonik und
Sedimentäre Geologie, Freie Universität Berlin, Fachbereich
Francis Albarède Ecole Normale Supérieure de Lyon, Lyon, France
Conel Michael O’Donel Alexander Department of Terrestrial Magnetism,
Carnegie Institution of Washington, NW Washington, DC, USA
Yann Alibert Space Research and Planetary Sciences, Physics Institute,
University of Bern, Bern, Swiss
Abigail Allwood Jet Propulsion Laboratory, Pasadena, CA, USA
Concepción Alonso Universidad Autonoma de Madrid, Madrid, Spain
Wladyslaw Altermann Department of Geology, University of Pretoria,
Pretoria, South Africa
Linda Amaral-Zettler Marine Biological Laboratory, Josephine Bay Paul
Center for Comparative Molecular Biology and Evolution, Woods Hole,
MA, USA
Ricardo Amils Departamento de Biologia Molecular, Universidad
Ariel D. Anbar School of Earth & Space Exploration and Department of
Chemistry & Biochemistry, Arizona State University, Tempe, AZ, USA
xxiii
xxiv Contributors
Luc André Department of Earth Sciences, Royal Museum of Central Africa,

Tervuren, Belgium
Philippe André Laboratoire AIM, IRFU/Service d’Astrophysique, CEA
Saclay, Gif-sur-Yvette, France
Ralf H. Anken German Aerospace Center (DLR), Institute of Aerospace
Medicine, Cologne, Germany
Josefa Anton Department of Physiology, Genetics and Microbiology,
University of Alicante, Alicante, Spain
Nicholas Arndt ISTerre, Université Grenoble Alpes, France
Andrew Aubrey NASA Jet Propulsion Laboratory, Pasadena, CA, USA
Jeffrey Bada Scripps Institution of Oceanography, La Jolla, CA, USA
Juan P. G. Ballesta Genome Dynamics and Function, Centro de Biologia
Molecular Severo Ochoa, Cantoblanco, Madrid, Spain
Nadia Balucani Dipartimento di Chimica, Università degli Studi di Perugia,
Perugia, Italy
Rory Barnes Astronomy Department, University of Washington, Seattle,
WA, USA
Maria Antonietta Barucci Observatoire de Paris, LESIA, Meudon Princi-
pal, Cedex, France
Gibor Basri Astronomy Department, MC 3411, University of California,
Berkeley, CA, USA
Ugo Bastolla Unidad de Bioinformática, Centro de Biologı́a Molecular
“Severo Ochoa,” CSIC-UAM, Madrid, Spain
Fabia Ursula Battistuzzi Oakland University, Rochester, MI, USA
Christa Baumstark-Khan German Aerospace Center (DLR), Institute of
Aerospace Medicine, Cologne, Germany
Andrey Bekker Department of Geology and Geophysics, Yale University,
New Haven, CT, USA
G. Fritz Benedict McDonald Observatory, The University of Texas, Austin,
TX, USA
Stefan Bengtson Department of Palaeozoology, The Swedish Museum of
Natural History, Stockholm, Sweden
Karim Benzerara Institut de Minéralogie et de Physique des Milieux
Condensés, UMR 7590, CNRS, Université Pierre et Marie Curie & Institut
de Physique du Globe de Paris, Paris, France
Jose Berenguer Centro de Biologı́a Molecular Severo Ochoa, UAM-CSIC,
Madrid, Spain
Contributors xxv
Sylvain Bernard Laboratoire de Minéralogie et de Cosmochimie du

Muséum (LMCM), Paris, France
Hugues Bersini IRIDIA, Université Libre de Bruxelles, Brussels, Belgium
Bruno Bézard LESIA, Observatoire de Paris, Meudon, France
Jean-Pierre Bibring Institut d’Astrophysique Spatiale, Université Paris

Sud, Orsay, France
John H. Black Department of Earth and Space Sciences, Chalmers Univer-

sity of Technology, Onsala Space Observatory, Onsala, Sweden
Donna Blackmond The Scripps Research Institute, La Jolla, CA, USA
Jeffrey Blanchard Biology Department, University of Massachusetts

Amherst, Amherst, MA, USA
Laurent Boiteau Institut des Biomolécules Max Mousseron, UMR5247

CNRS, University Montpellier-2, Montepellier Cedex, France
Tyler Bourke Square Kilometre Array Organisation, Macclesfield,

Cheshire, UK
Jessica C. Bowman School of Chemistry and Biochemistry, Georgia Insti-

tute of Technology, Atlanta, GA, USA
Samuel A. Bowring Department of Earth, Atmospheric and Planetary

Sciences, Massachusetts Institute of Technology, Building 54-1126,
Cambridge, MA, USA
Maud Boyet Université Blaise Pascal, Clermont-Ferrand, France
André Brack Centre de Biophysique Moléculaire CNRS, Orléans Cedex 2,

France
Robert Braun Square Kilometre Array Organisation, Macclesfield,

Cheshire, UK
Doris Breuer German Aerospace Center (DLR), Institute of Planetary

Research, Berlin, Germany
Carlos Briones Centro de Astrobiologı́a (CSIC/INTA), Consejo Superior de

Investigaciones Cientı́ficas, Madrid, Spain
Gilles Bruylants Engineering of Molecular NanoSystems, Universté Libre

de Bruxelles, Brussels, Belgium
Sergey Bulat NRC ‘Kurchatov institute’, Petersburg Nuclear Physics Insti-

tute, Leningrad Region, Gatchina, Russia
Vincent Busigny Institut de Physique du Globe de Paris, Paris, France
Gary R. Byerly Department of Geology & Geophysics, Louisiana State

University, Baton Rouge, LA, USA
xxvi Contributors
Michel Cabane LATMOS/IPSL B102/T45-46, Université Pierre et Marie

Curie UPMC-Paris 6, Paris, France
Jean Cadet Laboratoire Lésions des Acides Nucléiques, Institute
Nanosciences et Cryogénie/CEA, Grenoble, France
Michael P. Callahan Astrochemistry Laboratory, Code 691, NASA
Goddard Space Flight Center, Greenbelt, MD, USA
Jan Cami Department of Physics and Astronomy, The University of
Western Ontario, London, ON, Canada
SETI Institute, 189 Bernardo Avenue, Suite 100, Mountain View, CA, USA
Ian Campbell Research School of Earth Sciences, The Australian National
University, Canberra, ACT, Australia
Tammy Campbell Chemistry, The Scripps Research Institute, La Jolla,
CA, USA
Donald E. Canfield Institute of Biology, University of Southern Denmark,
Odense, Denmark
Marı́a Luz Cárdenas Unité de Bioénergétique et Ingénierie des Protéines,
Centre National de la Recherche Scientifique, Aix-Marseille Université,
Marseille Cedex 20, France
Damien Cardinal LOCEAN, Université Pierre & Marie Curie, Paris,
France
Leticia Carigi Instituto de Astronomı́a, Universidad Nacional Autónoma de
México, México, DF, Mexico
P. Brandon Carroll California Institute of Technology, Pasadena, CA, USA
Piergiorgio Casavecchia Dipartimento di Chimica, Università degli Studi
di Perugia, Perugia, Italy
Claude Catala 1LESIA, Observatoire de Paris, Meudon, France
Franco Cataldo Istituto Nazionale di Astrofisica – Osservatorio Astrofisico
di Catania, Catania, Italy
Actinium Chemical Research, Rome, Italy
David C. Catling Department of Earth and Space Sciences, University of
Washington, Seattle, WA, USA
Cecilia Ceccarelli Laboratoire d’Astrophysique de Grenoble (LAOG/
IPAG), Université J.Fourier de Grenoble, CNRS, Grenoble, France
Gilles Chabrier Centre de Recherche Astrophysiue de Lyon, Ecole
Normale Supérieure de Lyon, Lyon, France
John H. Chalmers Scripps Institute of Oceanography Geosciences
Research Division, University of California, San Diego, La Jolla, CA, USA
Contributors xxvii
Gregory Chambon Faculty of Arts and Humanities, University of Nantes,

Brest Cedex 3, France
Steven B. Charnley Solar System Exploration Division, Code 691,
Astrochemistry Laboratory, NASA Goddard Space Flight Center, Greenbelt,
MD, USA
Marc Chaussidon Institut de Physique du Globe de Paris (IPGP), Paris,
France
Glenn E. Ciolek New York Center for Astrobiology, Rensselaer Polytech-
nic Institute, Troy, NY, USA
Philippe Claeys Earth System Science, Vrije Universiteit Brussel, Brussels,
Belgium
Henderson James (Jim) Cleaves II Earth–Life Science Institute (ELSI),
Tokyo Institute of Technology, Meguro–ku, Tokyo, Japan
Institute for Advanced Study, Princeton, NJ, USA
Blue Marble Space Institute of Science, Washington, DC, USA
Center for Chemical Evolution, Georgia Institute of Technology, Atlanta,
GA, USA
Alain Coc Centre de Sciences Nucléaires et de Sciences de la Matière
(CSNSM) CNRS/IN2P3, Université Paris Sud 11, UMR 8609, Orsay, France
Charles S. Cockell Geomicrobiology Research Group, PSSRI, Open
University, Milton Keynes, UK
Catharine A. Conley NASA Headquarters, Washington, DC, USA
Joanna F. Corby University of Virginia, Charlottesville, VA, USA
Martin A. Cordiner The Goddard Center for Astrobiology, NASA Goddard
Space Flight Center, Greenbelt, MD, USA
Athel Cornish-Bowden Unité de Bioénergétique et Ingénierie des
Protéines, Centre National de la Recherche Scientifique, Aix-Marseille
Université, Marseille Cedex 20, France
Hervé Cottin Laboratoire Interuniversitaire des Systèmes Atmosphériques,
Université Paris Est-Créteil, Créteil, France
Athena Coustenis Laboratoire d’ Etudes Spatiales et d’ Instrumentation en
Astrophysique (LESIA), Observatoire de Paris, CNRS, UPMC Univ. Paris
06, Univ. Paris-Diderot, Meudon Cedex, France
Vanessa Cox Georgia Institute of Technology, Atlanta, GA, USA
Jacques Crovisier LESIA, Observatoire de Paris, Meudon, France
Alfonso F. Davila SETI Institute – NASA Ames Research Center MS 245-3,
Moffett Field, CA, USA
Thijs de Grauuw ALMA, Vitacura, Santiago, Chile
xxviii Contributors
Bradley De Gregorio Materials Science and Technology Division, U.S.

Naval Research Laboratory, Washington, DC, USA
Andrés de la Escosura Nanoscience and Molecular Materials Research
Group, Universidad Autónoma de Madrid, Madrid, Spain
Rafael R. de la Haba Department of Microbiology and Parasitology,
Faculty of Pharmacy, University of Sevilla, Sevilla, Spain
Jean-Pierre de Vera DLR, Institut f€ur Planetenforschung, Berlin, Germany
David Deamer Department of Chemistry, University of California, Santa
Cruz, Santa Cruz, CA, USA
Luis Delaye Departamento de Ingenierı́a Genética, CINVESTAV-Irapuato,
Irapuato, Gto, Mexico
René Demets ESTEC (HSF-USL), Noordwijk, The Netherlands
Didier Despois Laboratoire d’Astrophysique de Bordeaux, CNRS-
Universite de Bordeaux, France
Louis d’Hendecourt Institut d’Astrophysique Spatiale, Université
Paris-Sud 11, Orsay Cedex, France
Phil Diamond Square Kilometre Array Organisation, Macclesfield,
Cheshire, UK
Mark A. Ditzler NASA Ames Research Center, Moffett Field, CA, USA
Mark Dörr University of Southern Denmark, Odense M, Denmark
Thierry Douki Laboratoire Lésions des Acides Nucléiques, Institute
Nanosciences et Cryogénie/CEA, Grenoble, France
Nadja Drabon Stanford University, Stanford, CA, USA
Line Drube DLR Institute of Planetary Research, German Aerospace
Center (DLR), Berlin-Adlershof, Germany
Jean Duprat Centre de Sciences Nucléaires et de Sciences de la Matière
(CSNSM), Orsay, France
Claude D’Uston Géophysique Planétaire & Plasmas Spatiaux, Institut de
Recherche Astrophysique et Planétologique, Toulouse, France
Jason P. Dworkin NASA Goddard Space Flight Center, Astrochemistry
Laboratory, Greenbelt, MD, USA
Patrick Eggenberger Geneva Observatory, University of Geneva, Geneva,
Switzerland
Pascale Ehrenfreund Space Policy Institute, George Washington University,
Washington, DC, USA
Jennifer Eigenbrode NASA Goddard Space Flight Center, Greenbelt,
MD, USA
Contributors xxix
Sylvia Ekström Observatoire Astronomique de l’Université de Genève,

Faculté des Sciences, Université de Genève, Versoix, Switzerland
J. Cynan Ellis-Evans UK Arctic Office, Strategic Coordination Group,
British Antarctic Survey, Cambridge, UK
Josef Elster Faculty of Science, Centre for Polar Ecology, University of
South Bohemia, Ceske Budejovice, Czech Republic
Institute of Botany, Academy of Sciences of the Czech Republic, Trebon,
Czech Republic
Therese Encrenaz LESIA, Observatoire de Paris - Section de Meudon,
Meudon, France
Cecile Engrand Centre de Sciences Nucléaires et de Sciences de la Matière
(CSNSM), Orsay, France
Gözen Ertem National Institutes of Health, Bethesda, MD, USA
Alberto G. Fairén NASA Ames Research Center, Moffett Field, CA, USA
James Farquhar Department of Geology, University of Maryland, College
Park, MD, USA
Victor M. Fernández Institute of Catalysis, CSIC, Madrid, Spain
David C. Fernández-Remolar Centro de Astrobiologı́a (INTA-CSIC),
INTA, Torrejón de Ardoz, Spain
Fernando B. Figueiredo CITEUC, University of Coimbra, Coimbra,
Portugal
Ricardo Flores Instituto de Biologı́a Molecular y Celular de Plantas (UPV-
CSIC), Universidad Politécnica de Valencia – Consejo Superior de
Investigaciones Cientificas, Valencia, Spain
François Forget Institut Pierre Simon Laplace, Laboratoire de
Météorologie Dynamique, UMR 8539, Université Paris 6, Paris, France
Yves Fouquet Institut Français de Recherche pour l’Exploitation de la mer
(IFREMER), Issy-les-Moulineaux, France
Dionysis Foustoukos Geophysical Laboratory, Carnegie Institution of
Washington, Washington, DC, USA
Stephen Freeland Astrobiology Institute, University of Hawaii NASA,
Honolulu, HI, USA
Malcolm Fridlund Max-Planck-Institut f€ur Astronomie, Heidelberg,
Germany
Muriel Gargaud CNRS-Universite de Bordeaux, Laboratoire
d’Astrophysique de Bordeaux, Floirac, France
William Garnier Square Kilometre Array Organisation, Macclesfield,
Cheshire, UK
xxx Contributors
José Carlos Gaspar Institute of Geosciences, University of Brası́lia,

Brası́lia, DF, Brazil
Marı́a Gasset Consejo Superior de Investigaciones Cientı́ficas, Instituto
Quı́mica-Fı́sica Rocasolano, Madrid, Spain
Philipp Gast Asteroids and Comets, Institute of Planetary Research,
German Aerospace Center, Berlin, Germany
Eric Gaucher School of Biology, Georgia Institute of Technology, Atlanta,
GA, USA
B. Scott Gaudi Department of Astronomy, Ohio State University,
Columbus, OH, USA
Carlos Gershenson Instituto de Investigaciones en Matemáticas Aplicadas
y en Sistemas, Universidad Nacional Autónoma de México, DF, Mexico
Rosario Gil Institut Cavanilles de Biodiversitat i Biologia Evolutiva,
Universitat de València, Paterna (València), Spain
Felipe Gomez Centro de Astrobiologı́a (CSIC/INTA), Instituto Nacional de
Técnica Aeroespacial, Torrejón de Ardoz, Madrid, Spain
Aldo González Centro de Biologı́a Molecular, CBMSO Consejo Superior
de Investigaciones Cientificas Universidad Autónoma de Madrid, Madrid,
Spain
Elena González-Toril Laboratorio de Extremófilos, Centro de
Astrobiologı́a (INTA-CSIC), Torrejón de Ardoz, Madrid, Spain
Andrew Gould Department of Astronomy, Ohio State University, Colum-
bus, OH, USA
Matthieu Gounelle Laboratoire de Minéralogie et Cosmochimie du
Muséum (LMCM) MNHN USM 0205 - CNRS UMR 7202, Muséum National
d’Histoire Naturelle, Paris, France
Pierre-Henri Gouyon Département Systématique et Evolution, UMR 7138
CNRS-MNHN-UPMC-IRD, Muséum National d’Histoire Naturelle, Paris
Cedex 05, France
Felix M. Gradstein University of Oslo, Blindem, Oslo, Norway
Olivier Grasset University of Nantes, Nantes, France
Jimi Green Square Kilometre Array Organisation, Macclesfield, Cheshire,
UK
John Lee Grenfell German Aerospace Center (DLR), Berlin, Germany
Richard A. F. Grieve University of Western Ontario, London, ON, Canada
Elizabeth C. Griffith University of Colorado, Boulder, CO, USA
Roderich Groß Department of Automatic Control & Systems Engineering,
The University of Sheffield, Sheffield, UK
Contributors xxxi
udel Department of Astrophysics, University of Vienna, Vienna,

Manuel G€
Austria
Stephane Guillot LGCA, Universite de Grenoble, St Martin d’Hères,
France
Tristan Guillot Observatoire de la Côte d’Azur, Université de Nice-Sophia
Antipolis, CNRS, Nice, France
Weifu Guo Carnegie Institution of Washington, Washington, DC, USA
Nader Haghighipour Institute for Astronomy, University of Hawaii-
Gerhard Hahn Asteroids and Comets, DLR, Institute of Planetary
Salman Hameed Hampshire College, Amherst, MA, USA
Alan W. Harris DLR, Institute of Planetary Research, Berlin, Germany
Emma Hart School of Computing, Edinburgh Napier University,
Edinburgh, UK
Thomas H. P. Harvey Department of Earth Sciences, University of
Cambridge, Cambridge, UK
Ko Hashizume Department of Earth and Space Sciences, Osaka University,
Toyonaka, Osaka, Japan
Ernst Hauber Deutsches Zentrum f€ur Luft- und Raumfahrt (DLR) e.V.,
Institut f€ur Planetenforschung, Berlin, Germany
Rasmus Nielsen Haugaard Department of Earth and Atmospheric
Sciences, University of Alberta, Edmonton, AB, Canada
Robert Hazen Geophysical Laboratory, Carnegie Institution of Washington,
Washington, DC, USA
Jörn Helbert DLR, Institut f€ur Planetenforschung, Berlin, Germany
Ravit Helled Geophysical, Atmospheric and Planetary Sciences, Tel Aviv
University, Raymond and Beverly Sackler Faculty of Exact Sciences,
Tel Aviv, Israel
Ruth Hemmersbach German Aerospace Center (DLR), Institute of
Patrick Hennebelle Service d’Astrophysique, CEA, Saclay, Gif-sur Yvette,
France
Judith Herzfeld Brandeis University, Waltham, MA, USA
Christoph Heubeck Institut f€ur Geowissenschaften, Friedrich-Schiller-
Universität Jena, Jena, Germany
Ake Hjalmarson Chalmers University of Technology, Gothenburg, Sweden
xxxii Contributors
Pentti Hölttä Department of Geosciences and Geography, University of

Helsinki, Finland and Geological Survey of Finland, Espoo, Finland
Tori M. Hoehler Exobiology Branch, NASA Ames Research Center,
Mountain View, CA, USA
Paul Hoffman Department of Earth & Planetary Sciences, Harvard Univer-
sity, Cambridge, MA, USA
Harald Hoffmann DLR, Institute of Planetary Research, Berlin, Germany
Axel Hofmann Department of Geology, University of Johannesburg,
Auckland Park, Johannesburg, South Africa
Michiel R. Hogerheijde Leiden Observatory, Leiden University, Leiden,
The Netherlands
Martin Homann Institut f€ur Geologische Wissenschaften, Freie Universität
Berlin, Berlin, Germany
Gerda Horneck DLR German Aerospace Center, Institute of Aerospace
Medicine, Radiation Biology, Köln, Germany
David P. Horning Department of Molecular Biology, The Scripps Research
Institute, La Jolla, CA, USA
Nicholas V. Hud School of Chemistry and Biochemistry, Georgia Institute
of Technology, Atlanta, GA, USA
Elizabeth Humphreys ESO European Southern Observatory, Garching,
Germany
Susana Iglesias-Groth Instituto de Astrofisica de Canarias, La Laguna,
Tenerife, Spain
Heshan Grasshopper Illangkoon Department of Chemistry, University of
Florida, Gainesville, FL, USA
Eiichi Imai Nagaoka University of Technology, Nagaoka, Japan
William M. Irvine University of Massachusetts, Amherst, MA, USA
Akizumi Ishida Atmosphere and Ocean Research Institute, University of
Tokyo, Kashiwa, Chiba, Japan
Mathieu Isidro Square Kilometre Array Organisation, Macclesfield,
Cheshire, UK
Ralf Jaumann German Aerospace Center (DLR), Institute of Planetary
Emmanuelle J. Javaux Palaeobiogeology-Palaeobotany-Palaeopalynology,
Geology Department, Université de Liège, Liège, Belgium
Michel Jébrak Département des Sciences de la Terre et de l’Atmosphère,
Université du Québec à Montréal, Montreal, QC, Canada
Contributors xxxiii
Anders Johansen Lund University, Lund, Sweden

Natasha M. Johnson NASA Goddard Space Flight Center, Greenbelt,
MD, USA
Takeshi Kakegawa Graduate School of Science, Tohoku University,
Sendai, Japan
Paul Kalas Astronomy Department, University of California, Berkeley,
CA, USA
Lisa Kaltenegger Cornell University, Ithaca, NY, USA
Balz Samuel Kamber Trinity College, Dublin, Ireland
Inge Loes ten Kate Earth Sciences, Utrecht University, Utrecht,
The Netherlands
Michael J. Kaufman Department of Physics and Astronomy, San José State
University, San Jose, CA, USA
Kunio Kawamura Department of Human Environmental Studies,
Hiroshima Shudo University, Hiroshima, Japan
Yoko Kebukawa Department of Natural History Sciences, Hokkaido
University, Sapporo, Japan
Laura Kelly Ecogenomics of Interactions Lab, Nancy, France
Pierre Kervella LESIA, Observatoire de Paris, Meudon, France
Martin F. Kessler European Space Agency (ESA), European Space
Astronomy Centre (ESAC), Madrid, Spain
Daisuke Kiga Tokyo Institute of Technology, Tokyo, Japan
Eun-Kyong Kim Chemistry, The Scripps Research Institute, La Jolla,
CA, USA
Adrienne Kish Institut de Génétique et Microbiologie, Universite Paris-Sud
11, Orsay Cedex, France
Daniel Kitzmann University of Bern, Bern, Switzerland
David Klaus University of Colorado Boulder, Boulder, CO, USA
Thorsten Kleine Institut f€ur Planetologie, Westfälische Wilhelms-
Universität M€unster, M€unster, Germany
Kateryna Klochko Carnegie Institution of Washington, Washington,
DC, USA
Jörg Knollenberg Institute of Planetary Research, German Aerospace
Center, Berlin, Germany
Kensei Kobayashi Yokohama National University, Tokiwadai, Hodogaya-ku,
Yokohama, Japan
xxxiv Contributors
Kurt O. Konhauser Department of Earth and Atmospheric Sciences,

University of Alberta, Edmonton, AB, Canada
Akira Kouchi Institute of Low Temperature Science, Hokkaido University,

Kita-ku, Sapporo, Hokkaido, Japan
Ramanarayanan Krishnamurthy Chemistry, The Scripps Research

Institute, La Jolla, CA, USA
Edwin C. Krupp Griffith Observatory, Los Angeles, CA, USA
Marc Kuchner NASA Goddard Space Flight Center, Exoplanets and Stellar
Astrophysics Laboratory, Greenbelt, MD, USA
Ekkehard K€ uhrt Institute of Planetary Research, German Aerospace

Jana Kvı́derová Institute of Botany, Academy of Sciences of the Czech

Republic, Trebon, Czech Republic
Sun Kwok Faculty of Science, The University of Hong Kong, Hong Kong,
China
Jean-François Lambert Laboratoire de Réactivité de Surface, Université

Pierre et Marie Curie, Paris, France
Doron Lancet Weizmann Institute of Science, Rehovot, Israel
David W. Latham Harvard-Smithsonian Center for Astrophysics,

Cambridge, MA, USA
Amparo Latorre Institute Cavanilles for Biodiversity and Evolutionary

Biology, Universitat de Valencia, Valencia, Spain
Ester Lázaro Molecular Evolution Laboratory, Centro de Astrobiologı́a

(CSIC-INTA), Torrejón de Ardoz, Madrid, Spain
Antonio Lazcano Facultad de Ciencias, UNAM, Mexico, DF, Mexico
Stéphane Le Gars Centre François Viète, Université de Nantes, Nantes,

France
Michael Lebert Biology Department, Plant Ecophysiology, Friedrich-

Alexander-University Erlangen/Nuremberg, Erlangen, Germany
Laura M. Lechuga Nanobiosensors and Bioanalytical Applications Group,

Institut Català de Nanociència i Nanotecnologia (ICN2) CSIC and
CIBER-BBN, Barcelona, Spain
Guillaume Lecointre Département Systématique et Evolution, UMR 7138

CNRS-MNHN-UPMC-IRD, Muséum National d’Histoire Naturelle, Paris
Cedex 05, France
Emmanuel Lellouch Laboratoire d’Etudes Spatiales et d’Instrumentation

en Astrophysique (LESIA), Observatoire de Paris, Meudon, France
Contributors xxxv
Tom Lenaerts Département d’Informatique, Université Libre de Bruxelles,

Brussels, Belgium
Kevin Lepot Laboratoire d’Océanologie et de Géosciences, Université de

Lille, Villeneuve d’Ascq, France
Hugues Leroux Unité Matériaux et Transformations (UMET), University

Lille 1, Ronchin, Nord-Pas-de-Calais, France
Anny-Chantal Levasseur-Regourd UPMC University of Paris

6/LATMOS-IPSL, Paris, France
Richard Léveillé Natural Resource Sciences, McGill University, St. Anne

de Bellevue, Québec, Canada
Matthew Levy Michael F. Price Center, Albert Einstein College of

Medicine, Bronx, NY, USA
Purificación López-Garcı́a Unité d’Ecologie, Systématique et Evolution,

CNRS UMR8079 Université Paris-Sud 11, Paris, Orsay Cedex, France
Christophe Malaterre Institut d’Histoire et Philosophie des Sciences et

Techniques (IHPST), Université Paris 1-Panthéon Sorbonne, Paris, France
Physicalism Malaterre Institut d’Histoire et Philosophie des Sciences et

Techniques (IHPST), Université Paris 1-Panthéon Sorbonne, Paris, France
Irena Mamajanov School of Chemistry and Biochemistry, Georgia Insti-

tute of Technology, Atlanta, GA, USA
Rocco Mancinelli Bay Area Environmental Research Institute,

NASA Ames Research Institute, NASA Ames Research Center, Moffett
Field, CA, USA
Avi M. Mandell NASA Goddard Space Flight Center, Greenbelt, MD, USA
Susanna Manrubia Systems Biology Program, Centro Nacional de

Biotecnologı́a (CSIC), Madrid, Spain
Irma Marı́n Departamento de Biologı́a Molecular, Universidad Autónoma

de Madrid, Madrid, Spain
Lori Marino Emory Centre for Ethics, Emory University, Atlanta, GA, USA
Olivier La Marle Centre National d’Etudes Spatiales DSP/EU, Paris,

France
Mark S. Marley NASA Ames Research Center, Moffett Field, CA, USA
Jean-Emmanuel Martelat LST UMR5570, Université Claude Bernard

Lyon 1, St Martin d’Hères, Grenoble, France
Hervé Martin Laboratoire Magmas et Volcans, Université Blaise Pascal,

OPGC, CNRS, IRD, Clermont-Ferrand, France
xxxvi Contributors
Bernard Marty Institut Universitaire de France, Ecole Nationale

Supérieure de Géologie, Centre de Recherches Pétrographiques et
Géochimiques (CRPG), CNRS, Vandoeuvre les Nancy Cedex, France
Koichiro Matsuno Nagaoka University of Technology, Nagaoka, Japan
Thomas McCollom Laboratory for Atmospheric and Space Physics,
University of Colorado, Boulder, CO, USA
Francis McCubbin Institute of Meteoritics, University of New Mexico,
Albuquerque, NM, USA
Brett A. McGuire California Institute of Technology, Pasadena, CA, USA
Christopher P. McKay NASA Ames Research Center, Moffett Field,
CA, USA
Nicola McLoughlin Department of Earth Science and Centre for
Geobiology, University of Bergen, Bergen, Norway
Uwe J. Meierhenrich Institut de Chimie de Nice (ICN), University Nice-
Sophia Antipolis, Nice, France
H. Jay Melosh Departments of Earth, Atmospheric and Planetary Sciences,
Physics and Aerospace Engineering, Purdue University, West Lafayette,
IN, USA
Francesca Merlin CNRS UMR 8690 IHPST & Université Paris 1, Paris,
France
Allyssa Metzger Harvard University, Cambridge, MA, USA
François Mignard CNRS, Observatoire de la Côte d’Azur, University of
Nice Sophia-Antipolis, Nice, France
Stefanie N. Milam Astrochemistry Laboratory, NASA Goddard Space
Flight Center, Greenbelt, MD, USA
Thomas J. Millar Astrophysics Research Centre, School of Mathematics
and Physics, Queen’s University Belfast, Belfast, Antrim, UK
Vincent Minier CEA, Saclay, France
Shin Miyakawa Ribomic Inc., Minato-ku, Tokyo, Japan
A. M. Mloszewska Department of Earth and Atmospheric Sciences,
University of Alberta, Edmonton, AB, Canada
Robert Mochkovitch Institut d’Astrophysique de Paris, Paris, France
Ralf Moeller German Aerospace Center (DLR), Institute of Aerospace
Stephen Mojzsis University of Colorado, Boulder, CO, USA
Pierre-Alain Monnard FLinT center, Institute for Physics and Chemistry,
University of Southern Denmark, Odense M, Denmark
Contributors xxxvii
Francisco Montero Department of Biochemistry and Molecular Biology I,

Facultad de Ciencias Quı́micas, Universidad Complutense de Madrid,
Madrid, Spain
Thierry Montmerle Institut d’Astrophysique de Paris, CNRS/Université
Paris 6, Paris, France
Michel Morange Centre Cavaillès, USR 3308 CIRPHLES, Ecole normale
supérieure, Paris Cedex 05, France
Alessandro Morbidelli Observatoire de la Cote d’Azur, Nice, France
David Moreira Unité d’Ecologie, Systématique et Evolution CNRS
UMR8079, Université Paris-Sud 11, Paris, Orsay Cedex, France
Alvaro Moreno Departamento de Lógica y Filosofı́a de la Ciencia,
Universidad del Paı́s Vasco, San Sebastián, Spain
Miguel Moreno Centro de Astrobiologı́a, CSIC, Madrid, Spain
Harold Morowitz George Mason University, Fairfax, VA, USA
Stefano Mottola German Aerospace Center (DLR), Institute of Planetary
Denis J. P. Moura Ambassade de France en Italie, Ambascia di Francia,
Rome, Italy
Jean-François Moyen LMV-TL, Université Jean-Monnet, Saint-Etienne,
France
Armen Y. Mulkidjanian School of Physics, University of Osnabrueck,
Osnabrueck, Germany
Moscow State University, Moscow, Russia
uller I. Physikalisches Institut, Universität zu Köln, Köln,
Holger S. P. M€
Germany
Sami Nabhan Institute of Geological Sciences, Freie Universität Berlin,
Berlin, Germany
Kazumichi Nakagawa Graduate School of Human Development and
Environment, Kobe University, Nada, Kobe, Japan
Hiroshi Naraoka Department of Earth and Planetary Sciences, Kyushu
University, Fukuoka, Japan
Gopal Narayanan Five College Radio Astronomy Observatory, University
of Massachusetts, Amherst, MA, USA
Alicia Negrónk-Mendoza Instituto de Ciencias Nucleares, Universidad
Nacional Autónoma de México, México, Coyoacán, DF, Mexico
Gerhard Neukum Planetary Sciences and Remote Sensing, Institute of
Geological Sciences, Freie Universität Berlin, Berlin, Germany
xxxviii Contributors
Wayne L. Nicholson Space Life Sciences Laboratory, University of

Florida, Merritt Island, FL, USA
Space Life Sciences Laboratory, Kennedy Space Center, University of
Florida, Gainesville, FL, USA
Peter E. Nielsen The Panum Institute, ICMM, University of Copenhagen,
Copenhagen, Denmark
Nora Noffke Department of Ocean, Earth & Atmospheric Sciences, Old
Dominion University, Norfolk, VA, USA
Ann Nowé Vrije Universiteit Brussel, Brussels, Belgium
Joseph Andrew Nuth III Solar System Exploration Division, NASA’s
Goddard Space Flight Center, Greenbelt, MD, USA
Karin I. Öberg Harvard-Smithsonian Center for Astrophysics, Cambridge,
MA, USA
Shohei Ohara Carnegie Institution of Washington, Geophysical Labora-
tory, Washington, DC, USA
Hiroshi Ohmoto NASA Astrobiology Institute and Department of
Geosciences, The Pennsylvania State University, University Park, PA, USA
José Olivares Esztación Experimental del Zaidı́n. CSIC, Granada, Spain
Marc Ollivier Institut d’Astrophysique Spatiale, CNRS, Université de
Paris-Sud, Orsay, France
Hans Olofsson Department of Earth and Space Sciences, Chalmers Univer-
sity of Technology, Gothenburg, Sweden
Jonathan O’Neil Department of Earth and Environmental Sciences,
University of Ottawa, Ottawa, ON, Canada
Silvano Onofri Department of Ecological and Biological Sciences, Univer-
sity of Tuscia, Viterbo, Italy
Tullis C. Onstott Department of Geosciences, Princeton University,
Princeton, NJ, USA
Sijbren Otto Stratingh Institute for Chemistry, University of Groningen,
Groningen, The Netherlands
Corinna Panitz German Aerospace Center (DLR), Institute of Aerospace
Vı́ctor Parro Molecular Evolution Department, Centro de Astrobiologı́a
(INTA-CSIC), Torrejón de Ardoz, Madrid, Spain
Camille Partin Department of Geological Sciences, University of Manitoba,
Winnipeg, MB, Canada
Robert Pascal Institut des Biomolécules Max Mousseron CC1706,
Université de Montpellier II, Montpellier, France
Contributors xxxix
Matthew A. Pasek University of South Florida, Tampa, FL, USA

Mercedes Moreno Paz Molecular Evolution Department, Centro de
Astrobiologı́a (INTA-CSIC), Torrejón de Ardoz, Madrid, Spain
Ernesto Pecoits Department of Earth and Atmospheric Sciences, University
of Alberta, Edmonton, AB, Canada
Els Peeters Department of Physics and Astronomy, The University of
Western Ontario, London, ON, Canada
SETI Institute, 189 Bernardo Avenue, Suite 100, Mountain View, CA, USA
Ivanka Pelivan Institute of Planetary Research, German Aerospace Center,
Berlin, Germany
Juli Peretó Institut Cavanilles de Biodiversitat i Biologia Evolutiva,
Universitat de València, València, Spain
Jérôme Perez Applied Mathematics Laboratory, ENSTA ParisTech, Paris
Cedex 15, France
Jean-Robert Petit CNRS-UJF, Laboratoire de Glaciologie et Géophysique
de l’Environnement (LGGE), St Martin D’Heres, France
Pascal Philippot Equipe Géobiosphère Actuelle et Primitive, Institut de
Physique du Globe de Paris (IPGP), Paris, France
Ray Pierrehumbert Department of the Geophysical Sciences, University of
Chicago, Chicago, IL, USA
Göran L. Pilbratt ESA/ESTEC/SRE-S, European Space Agency (ESA),
Science Support Office, Noordwijk, The Netherlands
Samanta Pino Department of Biology and Biotechnologies “Charles
Darwin”, University of Rome “Sapienza”, Rome, Italy
Daniele L. Pinti GEOTOP Research Center for Geochemistry and
Geodynamics, Université du Québec à Montréal, Montréal, QC, Canada
Sandra Pizzarello Department of Chemistry & Biochemistry, Arizona State
University, Tempe, AZ, USA
Noah Planavsky Department of Geology and Geophysics, Yale University,
New Haven, CT, USA
Raphaël Plasson Department of Earth and Planetary Science, Harvard
University, Cambridge, MA, USA
Ana-Catalina Plesa German Aerospace Center (DLR), Institute of
Planetary Research, Berlin, Germany
Franck Poitrasson Geosciences Environnement Toulouse, CNRS,
Toulouse, France
Nikos Prantzos Institut d’Astrophysique de Paris, Paris, France
Lawrence Pratt Tulane University, New Orleans, LA, USA
xl Contributors
Daniel Prieur Université de Bretagne Occidentale (University of Western

Britanny), Brest, France
Institut Universitaire Européen de la Mer (IUEM), Technopôle Brest–Iroise,
Plouzané, France
José Cernicharo Quintanilla Department of Astrophysics, Laboratory of
Molecular Astrophysics, Iorrejón de Ardoz, Madrid, Spain
Ahmed Ragab Harvard Divinity School, Cambridge, MA, USA
Heike Rauer German Aerospace Center (DLR), Berlin, Germany
François Raulin Faculté des Sciences et Technologie, Université Paris Est
Créteil et Paris Diderot, LISA – UMR CNRS 7583, Creteil, France
Florence Raulin-Cerceau Maı̂tre de Conférences, Centre Alexandre Koyré
(UMR 8560-CNRS/EHESS/MNHN/CSI) Muséum National d’Histoire
Naturelle, Brunoy, France
Sean N. Raymond Laboratoire d’Astrophysique de Bordeaux, CNRS,
Universite de Bordeaux, France
Jacques Reisse Université Libre de Bruxelles, Brussels, Belgium
Anthony J. Remijan NRAO, Charlottesville, VA, USA
Petra Rettberg German Aerospace Center (DLR), Institute of Aerospace
Alonso Ricardo Ra Pharmaceuticals, Cambridge, MA, USA
Antonio J. Ricco NASA Ames Research Center, on Assignment from
Stanford University, Moffett Field, CA, USA
Wayne G. Roberge New York Center for Astrobiology, Rensselaer
Polytechnic Institute, Troy, NY, USA
François Robert Laboratoire de Minéralogie et Cosmochimie du Muséum
(LMCM), Muséum National d’Histoire Naturelle, UMR 7202 CNRS, Paris
Cedex 05, France
Michael P. Robertson Department of Molecular Biology MB42, The
Scripps Research Institute, La Jolla, CA, USA
Bernd Michael Rode Institute for General, Inorganic and Theoretical
Chemistry, Leopold-Franzens University, Universität Innsbruck, Innsbruck,
Austria
Francisco Rodriguez-Valera Microbiologia, Universidad Miguel
Hernandez, Campus San Juan, San Juan, Alicante, Spain
Françoise Roques Laboratoire d’Etudes Spatiales et d’Instrumentation en
Astrophysique (LESIA), Observatoire de Paris, Meudon, France
Minik T. Rosing Nordic Center for Earth’s Evolution, Natural History
Museum of Denmark, University of Copenhagen, Copenhagen, Denmark
Contributors xli
Ramon Rosselló-Móra IMEDEA (CSIC-UIB), Esporles, Mallorca,

Balearic Islands, Spain
Daniel Rouan LESIA, Observatoire Paris-Site de Meudon, Meudon, France
Kepa Ruiz-Mirazo Department of Logic and Philosophy of Science, FICE,

UPV-EHU, Biophysics Research Unit (CSIC – UPV/EHU), Donostia, San
Sebastián, Spain
Jan W. Sadownik Stratingh Institute for Chemistry, University of

Groningen, Groningen, The Netherlands
Nita Sahai Department of Polymer Science, University of Akron, Akron,

OH, USA
Cristina Sanchez-Porro Department of Microbiology and Parasitology,

Faculty of Pharmacy, University of Sevilla, Sevilla, Spain
Leopoldo G. Sancho Facultad de Farmacia Departamento de Biologia

Vegetal II, Universidad Complutense de Madrid, Madrid, Spain
José Luis Sanz Departamento de Biologı́a Molecular, Universidad

Pierre Savaton Université de Caen Basse-Normandie, Caen, France
Kevin Schindler Lowell Observatory, Flagstaff, AZ, USA
Karel Schulmann Ecole et Observatoire de Science de la Terre, Institute de

Physique de Globe, Université de Strasbourg, Strasbourg, France
Peter Schuster Institut f€ur Theoretische Chemie der Universität Wien,

Wien, Austria
Alan W. Schwartz Radboud University Nijmegen, Nijmegen, The

Netherlands
William G. Scott Department of Chemistry and Biochemistry, The Center

for the Molecular Biology of RNA, University of California at Santa Cruz,
Santa Cruz, CA, USA
Burckhard Seelig Department of Biochemistry, Molecular Biology and

Biophysics & BioTechnology Institute, University of Minnesota, St. Paul,
MN, USA
Antigona Segura Instituto de Ciencias Nucleares, Universidad Nacional

Autónoma de México, Mexico, DF, Mexico
Franck Selsis Laboratoire d’Astrophysique de Bordeaux, Universite de

Bordeaux, Floirac, France
CNRS, LAB, Floirac, France
Dmitry Semenov Max Planck Institute of Astronomy, Heidelberg,

Germany
xlii Contributors
Silke Severmann Institute of Marine & Coastal Sciences and Department of

Earth & Planetary Sciences, Rutgers University, New Brunswick, NJ, USA
M. A. Shea Air Force Research Laboratory (Emeritus), Bedford, MA, USA
Don F. Smart Air Force Research Laboratory (Emeritus), Bedford, MA, USA
Alexander Smirnov Department of Earth and Marine Science, Dowling

College, Oakdale, NY, USA
Ian W. M. Smith Chemistry Laboratory, University of Cambridge,

Cambridge, UK
Ronald L. Snell Department of Astronomy, 517 K Lederle Graduate

Research Center, University of Massachusetts, Amherst, MA, USA
Frank Sohl Deutsches Zentrum f€ur Luft- und Raumfahrt (DLR), Institut f€ur
Planetenforschung, Berlin, Germany
Alessandro Sozzetti Istituto Nazionale di Astrofisica (INAF) – Osservatorio

Astrofisico di Torino, Pino Torinese, Italy
Pietro Speroni di Fenizio CISUC, Department of Informatics Engineering,

University of Coimbra, Coimbra, Portugal
Tilman Spohn Deutsches Zentrum f€ur Luft- und Raumfahrt (DLR), Institut
f€
Greg Springsteen Furman University, Greenville, SC, USA
Steven W. Stahler Department of Astronomy, University of California,

Berkeley, CA, USA
Lucas J. Stal Department of Marine Microbiology, Royal Netherlands

Institute of Sea Research (NIOZ), Yerseke, The Netherlands
Vlada Stamenković Earth, Atmospheric and Planetary Sciences, Massachu-

setts Institute of Technology (MIT), Cambridge, MA, USA
Kenneth Mark Stedman Department of Biology, Center for Life in

Extreme Environments, Portland State University, Portland, OR, USA
Jennifer C. Stern Planetary Environments Laboratory, NASA Goddard

Barbara Stracke Deutsches Zentrum f€ur Luft- und Raumfahrt (DLR),

Institut f€
Harald Strauss Institut f€ur Geologie und Paläontologie, Westfälische

Wilhelms-Universität M€unster, M€unster, Germany
Kenichiro Sugitani Graduate School of Environmental Studies, Nagoya

University, Nagoya, Japan
Contributors xliii
Jun-Ichi Takahashi NTT Microsystem Integration Laboratories, Atsugi,

Japan
Olga Taran Chemistry and Chemical Biology, Harvard University,

Cambridge, MA, USA
Christophe Thomazo UMR CNRS 6282 Biogéosciences, Université de

Bourgogne, Dijon, France
Phil Thurston Laurentian University, Sudbury, ON, Canada
Simon Tillier Département Systématique et Evolution, UMR 7138 CNRS-

MNHN-UPMC-IRD, Muséum National d’Histoire Naturelle, Paris, France
Stéphane Tirard Centre François Viète d’Histoire des Sciences et des

Techniques EA 1161, Faculté des Sciences et des Techniques de Nantes,
Nantes, France
Daniela Tirsch German Aerospace Center DLR, Institute of Planetary

Dmitri Titov European Space Agency, Noordwijk, The Netherlands
Marco Tomassini Information Systems Department, University of

Lausanne, Lausanne, Switzerland
Carmen Tornow Institute of Planetary Research, German Aerospace

Melissa G. Trainer NASA Goddard Space Flight Center Code 699,

Greenbelt, MD, USA
Pascal Tremblin CEA, Saclay, France
Jorge L. Vago European Space Agency – ESA/ESTEC (SRE-SM),

Noordwijk, The Netherlands
Veronica Vaida University of Colorado, Boulder, CO, USA
Stephan van Gasselt Planetary Sciences and Remote Sensing, Institute of

Geological Sciences, Freie Universität Berlin, Berlin, Germany
Martin J. Van Kranendonk School of Biological, Earth and Environmen-

tal Sciences, University of New South Wales, Australia
Antonio Ventosa Department of Microbiology and Parasitology, Faculty of

Pharmacy, University of Sevilla, Sevilla, Spain
Enrique Viguera Genetics Department, Sciences Faculty, University of

Malaga, Malaga, Spain
Michel Viso CNES/DSP/SME, Vétérinaire/DVM, Astro/Exobiology, Paris

Cedex 1, France
xliv Contributors
unter von Kiedrowski Lehrstuhl f€ur Organische Chemie I, Ruhr-

G€
Universität Bochum, Bochum, NRW, Germany
Philip von Paris Laboratoire d’Astrophysique de Bordeaux, Universite de
Bordeaux, Floirac, France
Jeff Wagg Square Kilometre Array Organisation, Macclesfield, Cheshire,
UK
Roland J. Wagner German Aerospace Center (DLR), Institute of Planetary
Sara Imari Walker School of Earth and Space Exploration, Arizona State
University, Tempe, AZ, USA
Frances Westall Centre de Biophysique Moléculaire, CNRS, Orléans
Cedex 2, France
Hubert Whitechurch Ecole et Observatoire de Science de la Terre, Institute
de Physique de Globe, Université de Strasbourg, Strasbourg, France
Douglas Whittet New York Center for Astrobiology, Rensselaer Polytech-
nic Institute, Troy, NY, USA
Simon Wilde Department of Applied Geology, Curtin University of
Technology, Perth, Australia
Ian S. Williams Research School of Earth Sciences, ANU College of
Physical and Mathematical Sciences, Research School of Earth Sciences,
The Australian National University, Canberra, ACT, Australia
Loren Dean Williams School of Chemistry and Biochemistry, Georgia
Institute of Technology, Atlanta, GA, USA
Charles T. Wolfe Unit for History and Philosophy of Science, University of
Sydney, Sydney, NSW, Australia
Mark G. Wolfire Astronomy Department, University of Maryland, College
Park, MD, USA
Alexander Wolszczan Department of Astronomy & Astrophysics and Center
for Exoplanets & Habitable Worlds, The Pennsylvania State University,
University Park, PA, USA
Paul M. Woods Astrophysics Research Centre, School of Mathematics and
Physics, Queen’s University Belfast, Belfast, Antrim, UK
Kosei E. Yamaguchi Geochemical Laboratory, Department of Chemistry,
Toho University, Funabashi, Chiba, Japan
Masamichi Yamashita Institute of Space and Astronautical Science
(ISAS)/JAXA, Sagamihara, Kanagawa, Japan
Bruce Yardley School of Earth and Environment, University of Leeds,
Leeds, UK
Contributors xlv
Reika Yokochi Department of Geophysical Sciences, The University of

Department of Earth and Environmental Sciences, University of Illinois at
Philippe Zarka LESIA, Observatoire de Paris, CNRS, UPMC, Université

Paris Diderot, Meudon, France
Annie Zavagno CNRS, LAM (Laboratoire d’Astrophysique de Marseille)

UMR 7326, Aix Marseille Université, Marseille, France
Tanja Elsa Zegers Paleomagnetic Laboratory, Institute of Earth Sciences,

Utrecht University, Utrecht, CD, The Netherlands
Astrobiology by Discipline
Field - Astrophysics & Astrochemistry: Cosmic Ray, Ionization Rate

J. Cernicharo, M. Gargaud, W. Irvine, Cyanoethynyl Radical
D. Rouan Cyanogen Radical
Cyanomethanimine
Section - Astrochemistry: J. Cernicharo, Cyanopolyyne
W. Irvine, F. Salama Cyclopropenylidene
Diacetylene
4-Cyano-1,3-Butadiynyl Diazenylium
Absorption Spectroscopy Diffuse Interstellar Bands
Acetone Dihydroxyacetone
Acetylene Dimethyl Ether
Adsorption Dust Grain
Amino Radical Electron Attachment
Aminoacetonitrile Electron Dissociative Recombination
Ammonium (NH3D+) Electron Radiative Recombination
Amorphous Solid Elemental Depletion
Anion Eley-Rideal Mechanism
Apolar Molecule Ethanimine
Argonium Ethyl Cyanide
Benzene Ethyl Formate
Bimolecular Reaction Ethylene Glycol
Binding Energy Ethylene Oxide
Butadiynyl Radical Ethynyl Radical
C3H+ Extended Red Emission
Carbene Formamide
Carbodiimide Formyl Cation
Carbon Monosulfide Fullerane
Charge Transfer Fullerene
Chemical Bistability Gas-Grain Chemistry
Chemisorption Glycolaldehyde
Chlorine Hydrides in the Interstellar Medium Hydrogen Chloride
Circumstellar Chemistry Hydrogen Isocyanide
Clathrate Hydrate Hydrogenated Amorphous Carbon
CN- Hydroxyl Radical
Condensation Temperature Imidogen
# Springer-Verlag Berlin Heidelberg 2015

M. Gargaud et al. (eds.), Encyclopedia of Astrobiology,
DOI 10.1007/978-3-662-44185-5
2 Astrobiology by Discipline
Interstellar Chemical Processes Predissociation

Interstellar Ices Propyl Cyanide
Interstellar Molecule Propylene
Ion-Neutral Reaction Propynylidyne
IRC+10216 Proton Transfer
Isotopic Exchange Reaction Quenched Carbonaceous Composite
Isotopic Fractionation (Interstellar Medium) Radiative Attachment
Isotopolog Radical
Langevin Rate Coefficient Radiolysis
Langmuir-Hinshelwood Mechanism Reaction Rate Coefficient
Line Shielding Refractory Molecule
Metal Compounds in Circumstellar Envelopes Scattering
Methanethiol SgrB2
Methoxy Radical Silane
Methyl Acetate Silicon Monosulfide
Methyl Formate Silicon Monoxide
Methyl Radical Silicon Nitride
Methyl Triacetylene Sputtering
Methylene Star Dust
Methylidyne Sticking Coefficient
Methylidyne Cation Sulfur Hydrides in the Interstellar Medium
Molecular Abundances Sulfur Monoxide
Molecular Depletion Thioformaldehyde
Molecular Desorption TiO
Molecular Line Cooling TiO2
Molecular Line Map Unidentified Infrared Emission Bands
Molecular Line Survey Unimolecular Reaction
Molecules in Space UV Absorption Bump
Mutual Neutralization Vinyl Cyanide
Nanodiamond VY CMa
Neutral-Neutral Reaction Water in the Universe
Nitrogen Sulfide Water, Formation and Photodissociation
Nucleation of Dust Grains Water, Related Interstellar Radicals & Ions
Organic dust, synthesis by stars Water, Vibrational and Rotational Transitions
Organic Dust, Influence on the Origin of Life
Phosphaethyne
Phosphorus Monoxide Section - Astrophysics: General
Photochemistry Definitions: D. Rouan
Photodesorption
Photodetachment Ablation
Photodissociation Accretion Shock
Photoionization Activity, Magnetic
Photolysis Adaptive Optics
Physisorption Alignment of Dust Grains
Polar Molecule Angular Diameter
Polycyclic Aromatic Hydrocarbon Angular Momentum
Prasad-Tarafdar Mechanism Aphelion
Astrobiology by Discipline 3
Astrometry Hydrodynamic Flow

Atomic Fine Structure Cooling Hydrostatic Equilibrium
AU Imaging
Background Impact Parameter
Band Pass Inclination (Astronomy)
Blackbody Infrared Astronomy
Bolometer Interferometry
Bolometric Magnitude Interstellar Cloud
Bremsstrahlung Radiation Interstellar Dust
CCD Interstellar Medium
Celestial Equator Jeans Escape
Center of Mass Velocity Johnson UBV Bandpasses
Chandrasekhar’s Limit Lagrangian Points
Cirrus Cloud Light-Year
Coagulation, Interstellar Dust Grains Limb Darkening
Color Excess Limb, Astronomical
Color Index Line Emission
Column Density Line of Sight
Continuum Line Profile
Coordinate Systems Linewidth
Coronagraphy Local Standard of Rest
Cosmogony Luminosity
Declination Lyman Alpha
Dense Cloud Magnetic Field
Diffraction Magnitude
Diffuse Cloud Magnitude, Absolute
Diffuse Galactic Light Maser
Doppler Shift Mass Loss Rate
Dust Cloud, Interstellar Mean Free Path
Eccentricity Nadir
Ecliptic Noise
Effective Temperature Nulling Interferometry
Electromagnetic Radiation Occultation
Electromagnetic Spectrum Optical Depth
Emission Nebula Orbital Resonance
Emissivity Parallax
Ephemeris Parsec
Equation of State Photodissociation Region
Equinox Photon
Exozodiacal Light Photosphere
Extinction, Interstellar or Atmospheric Plasma
Flux, Radiative Polar Axis
Gas Giant Planet Precession
Gravitation Proper Motion
Grey Body Protoplanetary Nebula
Heavy Element Q (Orbital Parameter)
HII Region Radiative Processes
Radiative Transfer Asymptotic Giant Branch Star

Radio Astronomy Big Bang Nucleosynthesis
Red Rectangle Black Hole
Reddening, Interstellar CNO Cycle
Redshift Cosmochemistry
Reflection Nebula Diffusion
Right Ascension Drake Equation
Roche Limit Dwarf Star
Rotational Velocity Faint Young Sun Paradox
Semi Major Axis Fermi Paradox
Semi Minor Axis Galactic Habitable Zone
Shock, Interstellar Galaxy
Solar Constant Globular Cluster
Solar Luminosity Hertzsprung-Russell Diagram
Solar Mass High Mass Star
Solar Radius Horizontal Branch
Spectral Line Initial Mass Function
Spectrometer Isochrone
Spectroscopy Low Mass Star
Star Counts Main Sequence, Star
Suprathermal Mass-Luminosity Relation
Surface Gravity Metallicity
Synchrotron Radiation Milky Way
Telescope Neutron Star
Thermodynamical Chemical Equilibrium Nova
Time Series Nuclear Reaction
Titius-Bode Law Nuclear Stability
Translucent Interstellar Clouds Nucleosynthesis, explosive
Turbulence, Interstellar Nucleosynthesis, neutrino
Ultraviolet Radiation Nucleosynthesis, Stellar
UV Radiation Opacity
Vacuum Ultraviolet Open Cluster
Variability, Stellar Planetary Nebula
Vernal Point P-P Chains
Visible Light Pulsar
VLBI Red Dwarf
VLT Red Giant
XDR R-Process
X-rays (Stellar) Solar Neighborhood
Z Spallation Reaction
Zenith Spectral Type
S-process
Star
Section - Nucleosynthesis: N. Prantzos Stellar Evolution
Stellar Population
Abundances of Elements Stellar Pulsation
Asteroseismology Stellar Rotation
Stellar Yield Stellar Winds

Sun (and Young Sun) T Association
Supernova T Tauri Star
Supernova Remnant YY Orionis Star
Supernova Types
White Dwarf
Zero Age Main Sequence Field - Chemical Sciences: J. Cleaves
Section - Chemistry – General

Section - Stars – Formation and Definitions: K. Kobayashi, K. Bartik
Evolution: Ph. André
Acetaldehyde
Accretion, Stellar Acetic Acid
Ambipolar Diffusion Acetonitrile
Binary Stars, Young Activated Nucleotide
Bipolar Flow Adenine
Birthline Aerosols
Brown Dwarf Alcohol
Convection, Stellar Aliphatic Hydrocarbon
Debris Disk Alpha Rays
Dense Core Alteration
Fragmentation of Interstellar Clouds Amide
Free-Fall Time Amine
Gravitational Collapse, Stellar Amino Acid Precursors
Hot Core Aminoisobutyric Acid
Hot Corino Aminonitrile
Infrared Excess Amorphous Carbon
Initial Mass Function, Origin of Amphiphilicity
Interstellar Filaments Amphoteric Compounds
Larson’s Law Arginine
Lithium Absorption Asparagine
Magnetic Fields and Star Formation Aspartic Acid
Molecular Cloud Asymmetric Reaction, Absolute
OB Association Atmosphere, Organic Synthesis
Pillars ATP
Pre-main-sequence Star Beta Rays
Protobinary Star Black Smoker, Organic Chemistry
Protoplanetary Disk Carbon
Protostars Carbonyl
Protostellar Envelope Carboxylic Acid
Skumanich Law Catalyst
Spectral Classification of Embedded Stars Circular Dichroism
Spectral Veiling of Young Stars Clathrate
Star Formation, Observations Clay
Star Formation, Theory Complex Organic Molecules
Star Formation, Triggering Corona Discharge
Stellar Cluster Cosmic Ray in the Galaxy
Cyanoacetylene Insoluble Organic Matter

Cysteine Ionization Constant
Cystine Isoelectric Point
Cytosine Isoleucine
D/L-Ratio Isovaline
D-Amino Acids Kaolinite
Decarboxylation Ketose
Diamino Acid Kinetic Isotope Effect
Dicarboxylic Acid Lactic Acid
Diketopiperazine Leucine
Disulfide Bond Ligand
Endogenous Synthesis Lysine
Ester Mass Spectrometry
Ethanol Methane
Ether Methanol
Evolution, Chemical Methionine
Extraterrestrial Delivery of Organic Compounds Mildly Reducing Atmosphere
Extreme Ultraviolet Light Molecular Weight
Fischer Projection Monosaccharide
Formic Acid Montmorillonite
Fractionation Non-Protein Amino Acids
Free Amino Acid Nucleoside Phosphoimidazolide
Gamma Rays Oligomerization
GC/MS Oligonucleotide
Glutamic Acid Organic Molecule
Glutamine Organometallic
Glyceraldehyde Oxygen, Atomic
Glycerol pH
Glycolic Acid Phenylalanine
Guanine Photochemistry, Atmospheric
Halogen Polarized Electron
HCN Polymer Polarized Light and Homochirality
Heme Polymer
Heterocycle Polynucleotide
Histidine Polypeptide
Hoogsteen Pair Polysaccharide
Hydrocarbons Porphyrin
Hydrogen Prebiotic Chemistry
Hydrogen Cyanide Precursor
Hydrogen Sulfide Proline
Hydrolysis Propionaldehyde
Hydrophobicity Proteinoid Microsphere
Hydrothermal Reaction Proteins, Primary Structure
Hydrothermal Vent Origin of Life Models Proteins, Quaternary Structure
Hydroxy Acid Proteins, Secondary Structure
Hydroxyl Group Proteins, Tertiary Structure
Hypoxanthine Proton Irradiation
Ice Pyrolysis GC/MS
Quenching Activity
Radiochemistry Affinity Chromatography
Refractory Organic Polymer Affinity Constant
Ribonucleoside Alanine
Ribonucleotide Aldehyde
Sarcosine Aldose
Selenocysteine Alpha Helix
Serine Amino Acid
Shock Wave Amino Acid N-Carboxy Anhydride
Succinic Acid Amino Butyric Acid
Sulfur Ammonia
Supercritical Fluid Amphiphile
Svedberg Unit Aqueous Interfaces
Synchrotron Accelerator Aromatic Hydrocarbon
Synthetic Biology Arrhenius Plot
Tautomer Autocatalysis
Thermolysis Automaton, Chemical
Thiol Borate
Tholins Branching Ratio
Threonine B€ucherer-Bergs Synthesis
Thymine (T) Cahn Ingold Prelog Rules
Tryptophan Carbohydrate
Tyrosine Carbon Dioxide
Uracil (Ura) Carbon Monoxide
Urea Carbonaceous Chondrites, Organic Chemistry of
Valine Cell Models
Vesicle Chicken or Egg Problem
Volatile Chirality
VUV Chromatographic Co-elution
Water, Solvent of Life Chromatography
Watson-Crick Pairing Chromophore
Wave Number Combinatorial Library
Wavelength Combustion
Weak Bonds Composomes
Wobble Pair Concentration Gradients
XANES Covalent Bonds
X-rays (Organic Synthesis) Cryostat
Zwitterion Cyanamide
Cyanogen
Deamination
Section - Prebiotic Chemistry and Denaturation
Origins of Life: J. Cleaves, A. Pohorille Deoxyribose
Derivatization
Abiotic Photosynthesis Deuterium
Achiral Diastereomers
Acid Hydrolysis Dinitrogen
Activation Energy Dioxygen
Active Site Disproportionation
Dissolved Inorganic Carbon Equilibrium Membrane Potential

DNA Metabolism, Prebiotic
Double Helix Micelle
Electric Discharge Moiety
Electrophoresis Mole
Enantiomeric Excess Molecular Beacon
Enantiomers Molecular Recognition
Endergonic mRNA Display
Endothermic N-Carbamoyl-Amino Acid
Enthalpy Neutral Atmosphere
Entropy Nitrile
Evolution, Molecular Nitrogen
Exergonic Nucleic Acid Base
Exothermic Nucleic Acids
Fischer-Tropsch-Type Reaction Nucleon
Flow Reactor Nucleoside
Fluorescence Nucleotide
Fluorometry Nuclide
Fluorophore O/OREOS Nanosatellite
Formaldehyde Oligomer
Formose Reaction Oligopeptide
Free Energy Origin of Life
FRET Ornithine
Furanose Oxidizing Atmosphere
Gas Chromatography Permeability
Genetic Code Phase Transition
Globule, Nanoglobule Phosphine
Glycine Phosphite
Hapten Phosphoric Acid
HCNO Isomers PIXE
Heterotrophic Hypothesis PNA
Hexamethylenetetramine Polyoxymethylene
Homochirality Post-impact Plume
Homolysis Primordial Soup
HPLC p-RNA
Hydantoin Protein
Hydrogen Bond Protocell
Hydrophobic Effect Purine Bases
Hypercycle PVED
Infrared Spectroscopy Pyranose
Ion-Exchange Chromatography Pyrimidine Base
Isomer Pyrolysis
Isotopomer Pyrophosphate
L-Amino Acids Pyruvate
Ligase Quencher
Lipid Bilayer Racemic Mixture
Liquid Chromatography-Mass Spectrometry Racemization
Liquidus Radiation Pressure
Raman Spectroscopy Archean Tectonics

Redox Potential Banded Iron Formation
Ribose Barberton Greenstone Belt
Ribozyme Barberton Greenstone Belt, Sedimentology
Rice-Ramsperger-Kassel-Marcus Barberton Supergroup
RNA Baryte
RNA Ligase Basalt
RNA Replicase Ribozyme Black Smoker
RNA World Campbellrand-Malmani Platform, South Africa
Rotatory Power Canadian Precambrian Shield
Self-Assembly, Biological Carbonate on Mars
Self-Replication, Chemical Chert
Solidus Continental Crust
Spark Discharge Continents
Specific Activity Cool Early Earth
Stereochemistry Coonterunah Subgroup, Australia
Stereoisomers Craton
Steric Effect Crust
Strecker Synthesis Cryosphere
Sublimation Deccan Trapps
Substrate Diagenesis
Surface Plasmon Resonance Diamictite/Diamicton
Systems Chemistry Diapirism
Template-Directed Polymerization Dixon Island Formation, Western Australia
Thiocyanate Earth
Thioester Earth, Formation and Early Evolution
Trace Elements Earth, Surface Evolution
Transferase Evaporite
Triple Point Evaporites, Archean
Van Der Waals Forces Fennoscandia
Water Fig Tree Group
Fortescue Group
Ga
Field - Earth Sciences: D. Pinti Geological Timescale
Geothermal Gradient
Section - Archean Geology: Geothermobarometers
Ch. Heubeck Gondwana
Granite
Acasta Gneiss Greenland
Akilia Greenschist Facies
Amphibolite Facies Greenstone Belts
Anorthosite Hadean
Apex Basalt, Australia Huronian Glaciation
Apex Chert Hydrosphere
Archean Drilling Projects Hydrothermal Environments
Archean Environmental Conditions Igneous Rock
Archean Eon Isua Supracrustal Belt
Archean Mantle Jack Hills (Yilgarn, Western Australia)
Jaspilite Shield
Komatiite Silicate Minerals
Laurasia Snowball Earth
Lithosphere, Planetary Spherules
Ma Stratigraphy
Magma Subduction
Magnetic Anomaly Sulfate Minerals
Magnetic Pole Tides, Archean
Magnetite Tonalite-Trondhjemite-Granodiorite
Mantle Transvaal Supergroup, South Africa
Mantle Plume, Planetary Trapps
Metamorphic Rock Trondhjemite
Metamorphism Tumbiana Formation (Pilbara, Western
Metasediments Australia)
Metasomatism Turbidite
Mineral Volcaniclastic Sediment
Moho Volcano
Moodies Group Warrawoona Group
Moodies Group, Microbial Mats Weathering
MORB White Smoker
Mount McRae Shale Zircon
North Pole Dome (Pilbara, Western Australia)
Obduction
Oceanic Crust Section - Early Earth Geochemistry:
Oceans, Origin of F. Albarede
Onverwacht Group
Ophiolite Boron Isotopes
Paleomagnetism Carbonation
Paleosols Cerium, Anomalies of
Pangea Chalcophile Elements
Peridotite Decay Constant
Pilbara Craton Degassing
Pillow Lava Distillation, Rayleigh
Plate Tectonics Earth, Age of
Plate, Lithospheric Fischer-Tropsch Effects on Isotopic Fractionation
Precambrian Fluid Inclusions
Precambrian Oceans, Temperature of Fractionation, Mass Independent and Dependent
Proterozoic Eon Geochronology
Pyrite Graphite
Quartz Half-Life
Regolith, Terrestrial Hydrodynamic Escape
Rock Isochron
Rodinia Isotope
Sagduction Isotopic Fractionation (Planetary Process)
Sedimentary Rock Isotopic Ratio
Serpentine Late Veneer
Serpentinization Lithophile Elements
Shale Mantle, Oxidation of
Oxygen Fugacity Kerogen

Oxygen Isotopes Microbial Mats
Ozone Microbially Induced Sedimentary Structures
Phosphates Microfossils
Platinum Group Elements Microfossils, Analytical Techniques
Radioactivity Molecular Fossils
Rare Earth Elements Prokaryotes, Origin of
Siderophile Elements Pseudofossil
Sulfur Isotopes Rio Tinto
Shark Bay, Stromatolites of
Steranes, Rock Record
Section - Traces of Life: E. Javaux Stirling Range Biota
Stirling Range, Australia
Acid Maceration Strelley Pool Formation
Acritarch Stromatolites
Amoebae Sulfidic Oceans
Archean Traces of Life Syngenicity
Barberton Greenstone Belt, Traces of Early Life Transition Metals and Their Isotopes
Belcher Group, Microfossils
Biogenicity
Biomarkers Section - Geology and Geochemistry:
Biomarkers, Morphological General Definitions: D. Pinti
Biomineralization
Biopolymer Absolute and Relative Ages
Bioprecipitation Accretion
Biosignatures, Effect of Metamorphism Alunite
Biostabilization Amitsoq Gneisses
Bitumen Antarctica
Burgess Shale Biota Apex Chert, Microfossils
Cambrian Explosion Asthenosphere
Cap Carbonates Atacama Desert
Carbon Isotopes as a Geochemical Tracer Breccia
Carboxylic Acids, Geological Record of Bulk Silicate Earth
Chengjiang Biota, China Chemocline
Cyanobacteria, Diversity and Evolution of Chicxulub Crater
Dubiofossil Delta, Isotopic
Ediacaran Biota Deuterium/Hydrogen Ratio
Ediacaran Period Devon Island
Endogenicity Dharwar Craton
Eukaryotes, Appearance and Early Evolution of Earth’s Atmosphere, Origin and Evolution of
Exopolymers Ejecta
Fatty Acids, Geological Record of Extinct Radionuclides
Fossil Gabbro
Fossilization, Process of Geotherm
Gunflint Microbiota Geyser
Hopanes, Geological Record of Glaciation
Iron oxides, hydroxides and Oxy-hydroxides Goethite
Isoprenoids Great Oxygenation Event
Gunflint Formation Supercontinent

Hadean Mantle Tektite
Hematite Theia
Hydrogen Isotopes Thermonatrite
Hydrothermal Alteration Trona
Impact Degassing True Polar Wander, Theory of
Impact Melt Rock Ultramafic Rocks
Impactite Ultrastructure
Iridium Uraninite
Iron Isotopes Vostok, Subglacial Lake
Isotope Biosignatures Water, Delivery to Earth
Kaapvaal Craton, South Africa Weathering Profile
KREEP Yellowstone National Park, Natural Analogue
KT Boundary Site
Late Heavy Bombardment
Lomagundi Carbon Isotope Excursion
Mafic and Felsic Field - History and Philosophy of
Mantle Volatiles Science: S. Tirard
Mars Analogue Sites
Mass Extinctions Section - History and Philosophy of
Mid-Ocean Ridges Science: S. Tirard, F. Figueirido
Monomictic Breccia
Mud Volcano Abiogenesis
Natron Al-Andalus, Cosmological Ideas
Nitrogen Isotopes al-Bīrūnī, Abū Rayḥān
Noble Gases al-Tūsī, Nasir al-Dīn
Nuvvuagittuq (Porpoise Cove) Greenstone Belt Animalcules
Ocean, Chemical Evolution of Astrobiology
Olivine Baly’s Experiment
Oxygen Minimum Zone Bathybius Haeckelii
Oxygenation of the Earth’s Atmosphere Bernal’s Conception of Origins of Life
Permafrost Bruno, Giordano
Polimictic Breccia Buffon’s Conception of Origins of Life
Radiative Forcing Calvin’s Conception of Origins of Life
Radiogenic Isotopes Cassini, Giovani Domenico
Red Beds Cellular Theory, History of
Redox Zonation Chance and Randomness
Sanukitoid Co-evolution
Self-shielding Effects on Isotope Fractionation Comets, History of
Shocked Quartz Cosmic Background Radiation
Siderite Cosmogonia: Greece
Silicon Isotopes Cosmogonia: Mesopotamia
Soda Lakes Cosmogonia: Roma
Stable Isotopes Cuvier’s Conception of Origins of Life
Subglacial Environments Darwin’s Conception of Origins of Life
Suboxic De Duve, Christian
Sudbury Impact Structure de Maillet’s Conception of Origins of Life
Suevite Diderot’s Conception of Origins of Life
Dirac, Paul Urey’s Conception of Origins of Life

Earth’s Atmosphere, History of the Origins Vernadsky
Ecology, History of Vitalism
Enzymology: History of Woese, Carl
Evo-devo
Evolution of Species, Islamic Ideas
Fermi, Enrico Field - Institutions and Organizations:
Galilean Satellites W. Irvine, M. Viso
Gene, Selfish
Genetics, History of AEB
Geological Time Scale, History of ASA
Goldschmidt, Viktor Moritz ASI
Haeckel’s Conception of Origins of Life Bioastronomy (IAU Commission 51)
Haldane’s Conception of Origins of Life BNSC
Halley, Edmond CAB, Spain
Herschel, William CNES
Hoyle, Fred CNSA
Hubble, Edwin CONAE, Argentina
Huxley’s Conception on Origins of Life COSPAR
Ikhwan al-Safa CSA
Kant-Laplace Cosmogonic Hypothesis DLR, Germany
Kepler, Johannes DTU Space, Denmark
Lamarck’s Conception of Origins of Life EANA
Lemaı̂tre’s Theory of Expanding Universe (History) ECSS
Life in the Solar System (History) European Space Agency
Lowell, Percival IAF
Materialism IAU
Meteorites, History of IKI
Miller, Stanley ISO (Normative Organisation)
Monod’s Conception on the Origins of Life ISRO
Mythology ISSI
Native American Cosmology and Other Worlds ISSOL
Oparin’s Conception of Origins of Life JAXA
Origins of Life, History of JPL
Physicalism NAI
Planetary Theories and Cosmology, Islamic NASA
Theories NSO
Plank, Max Roskosmos
Plate Tectonics, History of SSB
Plurality of Worlds UK Space Agency
Protoplasmic Theory of Life
Radio Astronomy and Radio Telescopes, History of
Reductionism Field - Life Sciences: R. Amils
Sagan Carl
SETI, History of Section - Artificial Life: H. Bersini
Spectroscopy, History of
Spontaneous Generation, History of Artificial Chemistries
Todd, David Artificial Life
Biological Networks Electron Donor

Cellular Automata Electron Transport
Chemical Reaction Network Embden-Meyerhof-Parnas Pathway
Code Energy
Complexity Energy Conservation
Genetic Algorithms Entner-Doudoroff Pathway
Scale Free Networks Enzyme
Self Replication Exon
Self-Assembly Fermentation
Genome, Minimal
Gluconeogenesis
Section - Biochemistry: J. Pereto Glycolysis
Homeostasis
Aerobic Respiration Intron
Anabolism Life
Anaerobic Respiration Metabolism
Anoxygenic Photosynthesis Metabolism, Secondary
Antibody Mitochondrion
Anticodon NADH, NADPH
Assimilative Metabolism Nitrogen Cycle, Biological
ATP Synthase Nitrogen Fixation
ATPase Nucleus
Autopoiesis Photosynthesis
Autotroph Photosynthesis, Oxygenic
Autotrophy Photosynthetic Pigments
Bacteriochlorophyll Phototroph
Base Pair Primer
Bioenergetics Prion
Buffer Respiration
Calvin-Benson Cycle Restriction Enzyme
Catabolism RNA Polymerase
Cell Rubisco
Cell, Minimal Symbiosis
Chlorophylls Transcription
Chromosome Transduction
Citric Acid Cycle Transformation
Cloning Translation
Codon Wobble Hypothesis (Genetics)
Coenzyme
Cofactor
Cytochromes Section - Biology – General Definitions:
Cytoplasm F. Gomez
Diazotrophy
Dissimilative Metabolism Abiotic
DNA Polymerase Acidophile
Electrochemical Potential Aerobe
Electron Acceptor Algae
Electron Carrier Anaerobe
Biofilm Proton Pump

Carbon Cycle, Biological Psychrophile
Carbon Source Reducing Agent
Cell Wall Reduction
Colonization, Biological Sulfate Reducers
Cryptoendolithic Sulfur Cycle
Cyanobacteria Terrestrial Analog
Deep Biosphere Thermophile
Deep-Sea Microbiology Yeast
Deep-Subsurface Microbiology
Dormant State
Ecological Niche Section - Genetics and Evolution:
Ecosystem C. Briones
Endogenous
Endolithic Adaptation
Energy Sources Amplification (Genetics)
Environment Aptamer
Europa Analogues Aptasensor
Exogenous Biodiversity
Extreme Environment Bioinformatics
Extremophiles Cell Membrane
Habitat Cenancestor
Halophile Combinatorial Nucleic Acid Library
Halotolerance Common Ancestor
Heterotroph Conjugation
Hot Spring Microbiology DNA Sequencing
Hot Vent Microbiology Domain (Taxonomy)
Hypersaline Environment Endosymbiosis
Hyperthermophile Error Rate
Intelligence Evolution, Biological
Intelligence, Evolution of Evolution, In Vitro
Magnetosome Fidelity
Magnetotactic Bacteria Fitness
Mars Analogues Gene
Mesophile Gene Expression
Metabolic Diversity Genetic Map
Microorganism Genetics
Nitrification Genome
Organelle Genomics
Osmolite Genotype
Oxidation Homology
Oxygenase Hybridization
Peptidoglycan Last Universal Common Ancestor
Periplasm Lateral Gene Transfer
Piezophile Metagenome
Plankton Metatranscriptome
Proteobacteria Metavirome
Proton Motive Force Molecular Clock
Monophyletic Chemoautotroph
Mutagen Chemolithoautotroph
Mutagenesis Chemolithotroph
Mutant Chemoorganotroph
Mutation Chemotaxis
Natural Selection Chemotroph
Operon Chloroplast
Orthologous Gene Compatible Solute
Paralogous Gene Crenarchaeota
Phenetics Denitrification
Phenotype Eukarya
Phylogenetic Tree Euryarchaeota
Phylogeny Fungi
Phylotype Gaia Hypothesis
Phylum Genus
Plasmid Geomicrobiology
Polymerase Chain Reaction Glove Box
Proteome, Proteomics Gram-negative Bacteria
Quasispecies Gram-Positive Bacteria
Recombination Green Bacteria
Replication (Genetics) Hydrogenosomes
Ribosome Iron
Selection Iron Cycle
Sequence Korarchaeota
Sequence Analysis Lithotroph
Splicing Macronutrient
Systems Biology Membrane
Template Methanogens
Virion Methanotroph
Viroid Micronutrients
Virology Motility
Virus Multicellular Organisms
Nanoarchaeota
Nucleoid
Section - Life in Extreme Environments: Osmotic Pressure
R. Amils Outer Membrane
Oxic
Alkaliphile Peroxisome
Anoxic Photoautotroph
Antibiotic Pili
Archaea Planetary Ecosynthesis
Bacteria Ploidy
Bacterirhodopsin Prokaryote
Biogeochemical Cycles Protists
Biosensor Protoplast
Biosphere Quorum Sensing
Biotope Species
Carboxysomes, Structure and Function Species (Prokaryote)
Taq Polymerase
Taxonomy Field - Planetary Sciences and
Transport, Biological Exoplanets: T. Spohn
Unicellular Organisms
Water Activity Section - Exoplanetary Systems:
Xerophile N. Haghighipour
51 Pegasi B
Alpha Centauri Bb
Section - Microbiology in Space: Astrometric Orbit
G. Horneck Astrometric Planets
Barycenter
Aerobiology Beta Pictoris b
Apollo Mission Circumbinary Planet
Arrhenius Svante Circumprimary Planet
Biostack CoRoT 7b
Cosmic Rays in the Heliosphere Direct-Imaging, Planets
Desiccation Eclipse
DNA Damage Eta-Earth
DNA Repair Exomoon
Endospore Exoplanet, Detection and Characterization
Epilithic Exoplanets, Discovery
Gravitational Biology Exoplanets, Modeling Giant Planets’ Atmospheres
Hypolithic Fomalhaut b
HZE Particle Gamma Cephei
Ionizing Radiation, Biological Effects GJ 667C: First System with Multiple Super-
Lichens earths in Habitable Zone
Linear Energy Transfer Gliese 581, The Most Highly Debated Habitable
Lithopanspermia System
MEED Habitable Zone Around Binary Star Systems
Microgravity Habitable Zone in Binary Stars Systems
Ozone Layer Habitable Zone in Multi-star Systems
Photobiology HARPS
Planetary and Space Simulation Facilities HATNet
Radiation Biology HD 189733b
Radiation Dose HD 209458b
Solar Particle Events HIRES
Solar UV Radiation, Biological Effects Hot Jupiters
Space Biology Hot Neptunes
Space Environment HR 8799: The First Directly Imaged Multi-planet
Space Vacuum Effects System
Spallation Zone Kepler 10
Spore Kepler 11: Multiple Transiting Planet System
Sporulation Kepler 16b: First Circumbinary Planet
Survival Kepler 186f: First Earth-sized Planet in Habitable
UV Climate Zone
UV Radiation Dose Kepler 37b: A Moon-sized Planet
UV Radiation, Biological Effects Kepler 47: First Multi-circumbinary Planet System
Kepler 9: First Transiting System Confirmed by Asteroid Belt, Main

TTV Carbonaceous Chondrite
Keplerian Orbits Carbonate, Extraterrestrial
Light Travel Time Effect C-Asteroid
Microlensing Follow-Up Network Catena, Catenae
Microlensing Observations in Astrophysics Cavus, Cavi
Microlensing Planets Ceres
Mini-Neptunes Chaotic Region
Ocean Planet Chasma, Chasmata
OGLE-2005-BLG-390Lb Chassignites
OGLE-2006-BLG-109Lb,c Chassigny
Optical Gravitational Lensing Experiment Chondrite
Periastron Chondrule
Period Chronostratigraphy
Phase, Orbital CO2 Ice Cap (Mars)
Planet Detection: Transit Timing Variation CO2 Ice Clouds (Mars)
Planet detection; Eclipse Timing Variation Concretions (Mars)
Planets in Binary Star Systems Core, Planetary
Probing Lensing Anomalies Network Corona, Coronae
Pulsar Planets Cosmic Spherules
Radial Velocity Crater lakes (Mars)
Radial-Velocity Planets Crater, Impact
Rossiter-McLaughlin Effect Dark Streaks (Mars)
SETI Deimos
Spectroscopic Orbit Dichotomy, Planetary
Super-Earths Differentiation, Planetary
Transit Dust Devils
Transiting Planets Dwarf Planet
TrES Dynamo, Planetary
Eros Asteroid
Facula, Faculae
Section - Inner Solar System: A. Airo, Fossa, Fossae
B. Stracke Fumarole
Fusion Crust
6 Hebe Gaspra
Achondrite Gegenschein
Active Asteroid Gullies
Albedo Feature Habitability on Mars
ALH 84001 Heat Flow, Planetary
Amazonian Heat Transfer, Planetary
Annefrank Hesperian
Apollo Asteroid Hygiea
Apophis Asteroid Ida
Aquifer (Mars) Impact Basin
Arachnoid Interior Structure, Planetary
Areology Interplanetary Dust Particle
Asteroid Jarosite
JU3 Plume
Juno Polar Caps (Mars)
Labyrinthus, Labyrinthi Polar Layered Deposits (Mars)
Lacus Poynting-Robertson Drag
Landing Site Primordial Heat
Landslide (Mars) Psyche
Lava Tubes Radioactive Heating
Lenticula, Lenticulae Regio
Linea, Lineae Regolith, Planetary
Lingula, Lingulae Rheology, Planetary Interior
Lutetia Rille
Macula, Maculae Rima, Rimae
Magnetic Field, Planetary Rotation Planet
Mare, Maria RQ36
Mars Rupes, Rupēs
Mars Stratigraphy Satellite or Moon
Mathilde Selenology
Mensa/Mensae Serpentinization (Mars)
Mercury Shergottites
Meridiani (Mars) Shergotty
Meteor Slope Lineae, Recurrent
Meteoroid Slope streaks (Mars)
Mons, Montes Small Solar System Body
Moon, Origin of SNC Meteorites
Moon, The Sol
Nakhla Solar System, Inner
Nakhlites Solid-State Greenhouse Effect
Nanoparticle Space Weathering
Near-Earth Objects Stagnant Lid Convection
Nitrates on Mars Steins
Noachian Sulcus, Sulci
Obliquity and Obliquity Variations Sulfates, Extraterrestrial
Oceanus, Oceani Terra, Terrae
Olympus Mons Terrestrial Planet
Opaline Silica on Mars Tessera, Tesserae
Outflow Channels Tharsis
Pallas Tholus
Palus, Paludes Tides, Planetary
Panspermia Toutatis
Patera, Paterae Valles Marineris
Perchlorates on Mars Valley Networks
Phobos Vallis, Valles
Phosphates on Mars Vastitas, Vastitates
Phyllosilicates, Extraterrestrial Venus
Planet Venus Clouds
Planitia Vesta
Planum Zeolites
Section - Outer Solar System: Mimas

Th. Encrenaz Miranda
Neptune
Ariel Nereid
CAIs Oberon
Callisto Oort Cloud
Cassini Organic Refractory Matter
Cassini Division Parent Body
Centaurs (Asteroids) Parent Molecule, Comet
Charon Phoebe
Chiron Planetary Rings
Comet Pluto
Comet (Nucleus) Quaoar
Comet Borrelly Rhea
Comet Encke Saturn
Comet Giacobini-Zinner Sedna
Comet Hale-Bopp Solar System
Comet Halley Solar System, Outer
Comet Hartley 2 System Solar Formation, Chronology of
Comet Hyakutake Tethys
Comet Mc Naught Titan
Comet Shoemaker-Levy 9 Titania
Comet Shower Trans-Neptunian Object
Comet Tempel 1 Triton
Comet Wild 2 Trojans (Asteroids)
Cryovolcanism UltraCarbonaceous Antarctic Micrometeorites
Daughter Molecule, Comet Umbriel
Dione Uranus
Enceladus Water in the Solar System
Europa Zodiacal Light
Galileo Galilei
Ganymede
GEMs Section - Planetary and Exoplanetary
Giant Planets Atmospheres: J.L. Grenfell
Huygens
Iapetus Absorption Cross Section
Io Adiabatic Processes
Itokawa Asteroid Albedo
Jupiter AOGCM
Kuiper Belt Atmosphere, Escape
Lightcurve Atmosphere, Model 1D
Magnetosphere Atmosphere, Structure
Meteorite, Allende Atmosphere, Temperature Inversion
Meteorite, Murchison Atmospheric Habitability
Meteorite, Orgueil Bioindicator
Meteorites Biomarkers Atmospheric, Evolution Over Geo-
Micrometeorites logical Time
Biomarkers, Spectral Gas Drag

Clouds Giant Impact
Earth-like Atmosphere Gravitational Collapse, Planetary
Exoplanetary Atmospheric Retrieval Gravitational Focusing
False Negative Hill Radius/Sphere
False Positive Hill/Lagrange Stability
GCM Impact, Hit and Run
Greenhouse Effect Isolation Mass
Grey Gas Model Kozai Mechanism
Habitability of the Solar System Laplace Resonance
Habitability, Effect of Eccentricity Late-stage Accretion
Habitability, Effects of Stellar Irradiation Libration
Habitable Planet, Characterization Lindblad Resonance
Habitable Zone Magnetic Fields and Planetary Systems
Habitable Zone, Effect of Tidal Locking Formation
Hadley Cells Mean Motion Resonance
Latent Heat Meter-Size Catastrophe
Mie Scattering Nice Model
Non-Grey Gas Model: Real Gas Atmospheres Oligarchic Growth
Raman Scattering Orbit
Rayleigh Scattering Orbital Period
Scale Height Photoevaporation of Protoplanetary Disks
Stratosphere Planet Formation
Troposphere Planet V Hypothesis
Planetary Embryo
Planetary Evolution
Section - Planetary Formation and Planetary Migration
Dynamics: Y. Alibert, R. Helled Planetesimals
Protoplanetary Disk Dead Zone
Apsidal Angle Protoplanetary Disk Instability
Atmosphere, Primitive Envelope Protoplanetary Disk Midplane
Cassini State Protoplanetary Disk of Second Generation
Coagulation in Planetary Disks Protoplanetary Disk, Chemistry
Condensation Sequence Protosolar Nebula, Minimum Mass
Core Accretion, Model for Giant Planet Proto-sun Composition
Formation Q (Tidal Quality Factor)
Corotation Torque Q (Toomre Parameter)
Critical Core Mass (Giant Planet Formation) Q* (Specific Energy to Destroy an Object)
Disk Instability, Model for Giant Planet Radial Drift
Formation Runaway Gas Accretion
Dynamical Friction Runaway Growth
Dynamical Instability Secular Dynamics
Ejection, Hyperbolic Secular Resonance
Escape Velocity Shepherding
Feeding Zone Snow Line
Formation of Planetesimals - the Building Blocks Solar Nebula
of Planets Turbulence (Planetary Disks)
Viscosity Special Region (Mars)

Viscous Stirring Sporicide
Vortex, Vortices Sterile
Sterility Assurance Level
Sterilization
Field - Space Missions, Ground Terminal Sterilization Process
Facilities and Planetary Protection: Z-Value
M. Viso
Section - Space Missions and Ground
Section - Planetary Protection: M. Viso Facilities: M. Viso
Aerobic Mesophilic Bacterial Spores 67P/Churyumov-Gerasimenko

Aseptic Process ALMA
Assay Beagle 2
Biobarrier Biopan
Bioburden Cassini-Huygens Space Mission
Bioburden Controlled Environment Cheops
Bioburden Reduction COMET (Experiment)
Biodiversity (Planetary Protection) CoRoT Satellite
Biohazard Assessment Protocol Deep Impact
Biological Efficacy EPOXI Mission
Biological Indicator ERA
Biological Safety Level EURECA
Clean Room Exobiologie Experiment
Cleanliness ExoMars
Coleman-Sagan Equation Expose
Contamination, Probability Exposure Facilities
Depyrogenation Foton Capsule, Spacecraft
DHMR Gaia Mission
Disinfection Galileo Mission
D-Value Giotto Spacecraft
Encapsulated Bioburden Hayabusa Mission
Exposed Surface Bioburden Herschel Mission
Hard Landing Hipparcos
Heat Shock HST
HEPA Filters Huygens Probe
Impact, Probability Infrared Astronomical Satellite
Inactivation Infrared Space Observatory
Mated Bioburden International Space Station
Organic Material Inventory JUICE Mission
Outer Space Treaty JWST
Parametric Release Kepler Mission
Pasteurization Large Millimeter Telescope
Perennial Heat Source Long Duration Exposure Facility
Planetary Protection Mars 2020
Planetary Protection Category Mars Express
Quarantine Mars Global Surveyor
Sample Receiving Facility Mars Odyssey
Mars Orbiter Mission Submillimeter Wave Astronomy Satellite

Mars Pathfinder TESS
Mars Reconnaissance Orbiter TPF/Darwin
Mars Sample Return Mission Ulysses Mission
Mars Science Laboratory Vega 1 and 2 Spacecraft
MER, Spirit and Opportunity (Mars) Viking
Odin Voyager, Spacecraft
Philae Lander WASP
Phobos-Grunt Yinghuo-1
Phoenix
Pioneer Spacecraft
PLATO 2.0 Satellite Field - Data Tables: M. Gargaud
Rosetta Spacecraft
SIM Astronomical Data
Spitzer Space Telescope Chemical and Biological Data
Square Kilometre Array Chronological History of Life on Earth
Stardust Mission General Data
STONE Geological Data
A
AAN Abiogenic Photosynthesis
▶ Aminoacetonitrile ▶ Abiotic Photosynthesis
Abiogenesis
Abiotic
Stéphane Tirard
Centre François Viète d’Histoire des David C. Fernández-Remolar
Sciences et des Techniques EA 1161, Faculté Centro de Astrobiologı́a (INTA-CSIC), INTA,
des Sciences et des Techniques de Nantes, Torrejón de Ardoz, Spain
Nantes, France
Definition
Definition
Abiotic refers to the physical and chemical pro-
Thomas Huxley (1825–1895) used the term abio- cesses that take place in natural environments but
genesis in an important text published in 1870. are driven by mechanisms that do not involve any
He strictly made the difference between sponta- biological activity. Although major physical and
neous generation, which he did not accept, and chemical cycles on Earth can hardly escape the
the possibility of the evolution of matter from activity of the biosphere, some processes do not
inert to living, without any influence of life. depend on biological activities. For example, this
Since the end of the nineteenth century, is the case of the formation of hydrothermal
evolutive abiogenesis means increasing complexity deposits that are based on redox, volatile fugac-
and evolution of matter from inert to living state in ity, and high thermal conditions. Some abiotic
the abiotic context of evolution of primitive Earth. processes are involved in the production of sur-
face oxidants through photochemical reactions in
planet atmospheres as has been proposed to
See Also explain the presence of perchlorates on Mars.
Paradoxically, different abiotic pathways
▶ Darwin’s Conception of the Origins of Life (thermal, radiolytic, or photochemical) create
▶ Huxley’s Conception on Origins of Life the chemical disequilibrium which is strictly nec-
▶ Origin of Life essary to fuel physical and chemical cycles on
DOI 10.1007/978-3-662-44185-5
26 Abiotic Photosynthesis
planets depleted of life. Some abiotic pathways History

have likely been essential to originate the
primary biochemical machinery that drove the In September 1912, Benjamin Moore suggested
emergence of life on Earth. In this sense, most at a discussion on the origin of life, held by the
abiotic processes support the production of the joint sections of Zoology and Physiology of the
compounds and chemical disequilibrium essen- British Association for the Advancement of Sci-
tial for a region of the universe to become ence, that “the first step towards the origin of life
habitable. must have been the synthesis of organic matter
from inorganic by the agency of inorganic col-
loids acting as transformers or catalysts for radi-
ant solar energy” (Moore and Webster 1913).
See Also
▶ Hydrothermal Environments
Overview
▶ Origin of Life
▶ Photochemistry
In spite of Haldane’s well-known idea that UV
▶ Prebiotic Chemistry
light may have served as a driving force for
formation of the first viruslike organisms
(Haldane 1929), the idea of directly driving abio-
genesis by solar energy had not won much sup-
Abiotic Photosynthesis port at that time, despite the fact that the Sun is by
far the most powerful energy source on Earth.
Armen Y. Mulkidjanian The limited acceptance of the idea was partly
School of Physics, University of Osnabrueck, due to the low quantum yield of abiotic photo-
Osnabrueck, Germany synthetic reactions and the poor reproducibility
Moscow State University, Moscow, Russia of experimental results. Abiotic photoproduction
of hydrogen, in the presence of ions of divalent
iron, has been observed (Mauzerall et al. 1993). It
Keywords has been shown atmospheric photochemistry can
produce aldehydes from CO (Bar-Nun and Chang
Bacterial photosynthesis; Carbon fixation; Pho- 1983). Only in the 1980s, were robust procedures
tochemistry; Semiconductors; Anoxic geother- of producing colloidal nanoparticles of
mal fields photoactive semiconductors, such as zinc sulfide
(ZnS) or cadmium sulfide (CdS), developed
(Henglein 1984). These particles (see Fig. 1),
due to their high surface-to-volume ratio, pro-
Synonyms
vided experimental systems in which the photo-
reduction of CO2 to diverse organic compounds
Abiogenic photosynthesis; Prebiotic
could be studied. The photoreduction proceeded
photosynthesis
with high and reproducible quantum yield (up to
80 % for CO2 reduction to formate at the surface
of colloidal ZnS particles (Henglein 1984)).
Definition Recent studies have demonstrated high-yielding
ZnS- and MnS-mediated photosynthesis under
Abiotic or abiogenic photosynthesis is the syn- simulated primeval conditions (Zhang
thesis of organic compounds with the aid of radi- et al. 2004, 2007; Guzman and Martin 2009). In
ant energy and various inorganic or organic the modern oceans, ZnS and MnS are found at the
catalysts. sites of geothermal activity, where minute
Abiotic Photosynthesis 27
E, V (redox potential)
A
Surface
-2 electron trap
Conduction band
CO2 + 2H+
0
H2S S2-
HCOOH
S-
S + 2H+
2 UV light
Valence band
Abiotic Photosynthesis, Fig. 1 Abiogenic photosyn- surface, where they can be picked up by appropriate
thesis on the primordial Earth. Left panel: light-induced acceptors, e.g., molecules of CO2. The residual electron
reactions in a ZnS particle combined with an energy vacancies (holes) are initially reduced by the S2 ions of
diagram. The absorption of a UV quantum by a minute the crystal, which then eventually can be replenished by
crystal of ZnS, an n-type semiconductor, leads to the external electron donors, e.g., H2S (cf with the mechanism
separation of electric charges and to the transition of the of anoxygenic photosynthesis). Right panel: the precipi-
excited electrons into the conducting zone. The electrons tation of ZnS particles (gray dots) around a Hadean con-
can migrate inside the crystal until they are trapped at the tinental hot spring (Figure from Mulkidjanian 2009)
particles of these minerals continuously precipi- fumaroles. The development of the first life forms
tate around hot, deep-sea hydrothermal vents; within photosynthesizing, ZnS-containing pre-
thereby, particles of ZnS and MnS, slowly precipitates at such anoxic geothermal fields, where
cipitating sulfides, make rings around black Zn2+ ions would be continuously released as
throats of such vents that are covered by promptly by-products of abiogenic photosynthesis, might
precipited particles of FeS (Tivey 2007). On the explain cellular enrichments in Zn2+, the equilib-
primordial Earth, hot metal-enriched geothermal rium concentration of which in the primordial
fluids and vapor may have discharged to the sur- ocean should have been extremely low
face of the first continents, so that particles of ZnS (Mulkidjanian and Galperin 2009; Mulkidjanian
and MnS could have precipitated within regions et al. 2012). Several proteins shared by all extant
exposed to solar radiation (Mulkidjanian 2009). organisms and believed to form the core of the
These sulfide minerals could have been present in last universal common ancestor (LUCA) are par-
shallow waters (Guzman and Martin 2009) and ticularly enriched in Zn and Mn; this may also
should have precipitated around continental ther- support the notion of a role for abiogenic photo-
mal springs (Mulkidjanian 2009). Since Zn2+ synthesis in the earliest stages of evolution
ions are much more volatile than Fe2+ ions, the (Mulkidjanian and Galperin 2009; Mulkidjanian
vapor of continental geothermal systems would et al. 2012). Since these ubiquitous proteins are
be particularly enriched in ZnS (Mulkidjanian depleted in iron, it remains to be established
et al. 2012). On the primordial Earth, ZnS could whether and to what extent iron (II), the predom-
not be oxidized by atmospheric oxygen, so that inant transition metal in geothermal exhalations,
photosynthesizing and habitable rings may have was involved in abiogenic photosynthesis. It has
persisted around terrestrial thermal springs and also been shown that titanium dioxide particles
28 Ablation
can drive photosynthetic organic chemistry Moore B, Webster TA (1913) Synthesis by sunlight
inside cell membrane-like vesicles (Summers in relationship to the origin of life. Synthesis of
formaldehyde from carbon dioxide and water by
et al. 2009). Titanium dioxide (both rutile and inorganic colloids acting as transformers
anatase) particles could have been formed by of light energy. Proc R Soc Lond B Biol Sci
precipitation or released (directly or from alter- 87:163–176
ation of other titanium minerals) by weathering. Mulkidjanian AY (2009) On the origin of life in the Zinc
world: 1. Photosynthetic, porous edifices built of
This energy transduction could have provided hydrothermally precipitated zinc sulfide (ZnS) as cra-
pathways to new compounds in a prebiotic sys- dles of life on Earth. Biol Direct 4:26
tem or support early biochemical reactions. Mulkidjanian AY, Galperin MY (2009) On the
origin of life in the Zinc world. 2. Validation of
the hypothesis on the photosynthesizing zinc
sulfide edifices as cradles of life on Earth. Biol
Direct 4:27
See Also Mulkidjanian AY, Bychkov AY, Dibrova DV, Galperin
MY, Koonin EV (2012) Origin of first cells at terres-
▶ Anoxygenic Photosynthesis trial, anoxic geothermal fields. Proc Natl Acad Sci
U S A 109:E821–E830
▶ Black Smoker Summers DP, Noveron J, Basa RCB (2009) Energy trans-
▶ Carbon Dioxide duction inside of amphiphilic vesicles: encapsulation
▶ Charge Transfer of photochemically active semiconducting particles.
▶ Earth’s Atmosphere, Origin and Evolution of Orig Life Evol Biosph 39:127–140
Tivey MK (2007) Generation of seafloor hydrothermal
▶ Electron Acceptor vent fluids and associated mineral deposits. Oceanog-
▶ Electron Donor raphy 20(1):50–65
▶ Energy Sources Zhang XV, Martin ST, Friend CM, Schoonen MAA, Hol-
▶ Extreme Ultraviolet Light land HD (2004) Mineral-assisted pathways in prebiotic
synthesis: photoelectrochemical reduction of carbon
▶ Formic Acid (+IV) by manganese sulfide. J Am Chem Soc
▶ Haldane’s Conception of Origins of Life 126(36):11247–11253
▶ Hot Spring Microbiology Zhang XV, Ellery SP, Friend CM, Holland HD, Michel
▶ Hydrothermal Vent Origin of Life Models FM, Schoonen MAA, Martin ST (2007) Photodriven
reduction and oxidation reactions on colloidal
▶ Iron semiconductor particles: implications for prebiotic
▶ LUCA synthesis. J Photochem Photobiol A Chem 185(2–3):
▶ Origin of Life 301–311
▶ Photochemistry
▶ Photosynthesis
▶ Transition Metals and Their Isotopes
▶ UV Radiation
▶ White Smoker Ablation
Daniel Rouan
References and Further Reading LESIA, Observatoire Paris-Site de Meudon,
Meudon, France
Guzman MI, Martin ST (2009) Prebiotic metabolism:
production by mineral photoelectrochemistry of
alpha-ketocarboxylic acids in the reductive tricarbox-
ylic acid cycle. Astrobiology 9(9):833–842
Haldane JBS (1929) The origin of life. Rationalist annual. Definition
Watts & Co, London, pp 3–10
Henglein A (1984) Catalysis of photochemical reactions Ablation is the erosion of the surface of a solid
by colloidal semiconductors. Pure Appl Chem object in a flow (e.g., during the entrance of an
56(9):1215–1224
Mauzerall D, Borowska Z, Zielinski I (1993) Photo and object into the atmosphere) through some physi-
thermal reactions of ferrous hydroxide. Orig Life cal process, such as formation of a ▶ fusion crust,
Evol Biosph 23(2):105–114 vaporization, or friction.
Absorption Spectroscopy 29
scattered into other angles. The rate at which

Absolute and Relative Ages energy is taken out of the beam by absorption
and scattering can be characterized in terms of A
Daniele L. Pinti coefficients with dimensions of area, which are
GEOTOP Research Center for Geochemistry and known as cross sections. The term absorption
Geodynamics, Université du Québec à Montréal, cross section is often used to include both the
Montréal, QC, Canada portion due to scattering and that due to true
absorption (loss of the photon into another form
of energy, such as heat). For atmospheric gases,
Definition this total absorption cross section is defined by
the Beer’s law expression:
In ▶ geochronology, the absolute age of a rock is
the age obtained from the measurement of spon- I ¼ I 0 expðsnlÞ
taneous decay of radioactive nuclides contained
in the rocks or its constituent minerals. It differs where I0 and I are the incident and transmitted
from the relative age. This latter is the age of a light intensities, respectively, s is the absorption
rock obtained from time and space relations cross section (cm2 molecule1), n is the molecu-
between rock formations, giving only qualitative lar density, and l is the pathlength in cm.
or semiquantitative information on the period
when the rock formed.
Absorption Spectroscopy
See Also
Steven B. Charnley
▶ Earth, Age of
Solar System Exploration Division, Code
▶ Geochronology
691, Astrochemistry Laboratory, NASA Goddard
▶ Radioactivity
Definition
Absorption Cross Section
In absorption ▶ spectroscopy, the spectral fea-
Lisa Kaltenegger tures of interest appear in absorption with respect
Cornell University, Ithaca, NY, USA to a background continuous spectrum. In the
interstellar medium, the background continuum
may be supplied by a radiation source, such as a
Keywords star, located behind the region of interest. The
absorbing material may be either in the gas or
Absorption the solid phase (e.g., interstellar dust or ices).
Solid state features are much broader than atomic
or molecular absorptions and are consequently
Definition more difficult to assign to a specific carrier.
Much of the solar (Fraunhofer) spectrum is seen
When a parallel, monochromatic beam of light in absorption, as the outer cooler layers of the
traveling in some specific direction encounters a solar atmosphere absorb radiation from the
medium of finite extent, a certain amount of the deeper photosphere. Spectral lines in planetary
flux will be absorbed and a certain amount will be atmospheres are typically seen in absorption,
30 Abundances of Elements
against the continuous thermal spectrum from the are key ingredients for understanding their origin
planetary or satellite surface. and evolution.
Overview
History
The composition of remote objects (the Sun,
The first person to notice a number of dark fea-
▶ stars, interstellar gas, and galaxies) is deter-
tures in the solar spectrum was the English chem-
mined through spectroscopy, which usually
ist William Wollaston in 1802. This absorption
allows the determination of elemental abun-
spectrum was first systematically investigated by
dances; in rare cases, particularly for interstellar
Joseph von Fraunhofer, starting in 1814, and the
clouds, some isotopic abundances may be deter-
spectral features are now known as Fraunhofer
mined in those objects. For Earth, lunar, and
lines.
meteoritic samples, nuclear mass spectroscopy
allows precise determination of most isotopic
abundances; this is also the case for cosmic
See Also rays, albeit only for the most abundant nuclides
at present. Hydrogen (H) being the most abun-
▶ Spectroscopy dant element in the Universe, spectroscopists
express the abundance of element i as the number
ratio of its nuclei with respect to those of H:
ni = Ni /NH, and they use a scale where
log(NH) = 12. In the meteoritics community,
Abundances of Elements
the silicon scale of log(NSi) = 6 is used. Theore-
ticians use the mass fraction Xi = Ni Ai / Nj Aj,
Nikos Prantzos
where Aj is the mass number of nuclide j; obvi-
Institut d’Astrophysique de Paris, Paris, France
ously, Xi = 1. Conversion of mass fractions to
abundances by number requires use of the quan-
tity Yi = Xi /Ai called the mole fraction (notice
Keywords
that Yi 6¼ 1).
According to our current understanding, the
Chemical composition; Nucleosynthesis;
material of the proto-solar nebula had a remark-
Nuclide
ably homogeneous composition, as a result of
high temperatures (which caused the melting of
nearly all the dust grains) and thorough mixing.
Definition This composition characterizes the present-day
surface layers of the Sun, which remain unaf-
The relative amount (or fraction) of a given fected by nuclear reactions occurring in the
nuclide in a sample of matter is called the abun- solar interior (with a few exceptions, e.g., the
dance of that nuclide. It can be expressed either in fragile D and Li). Furthermore, after various
absolute terms (i.e., with respect to the total physicochemical effects are taken into account,
amount of matter in the sample) or in relative it appears that the elemental composition of the
terms (with respect to the amount of some key Earth and meteorites matches extremely well
element, e.g., the most abundant one, in the sam- with the solar photospheric composition. The
ple). Similarities and differences in the elemental composition of stars in the Milky Way presents
and isotopic composition of ▶ stars and galaxies both striking similarities and considerable
Abundances of Elements 31
11
H
10 He A
9
8
O
C
7
Log (Abundance) [Si = 6]
Ne
Mg Si
N Fe
6 S
Ar
5 Ca Ni
Al
Na Cr
4
P Ti Mn
Cl K Zn
3 Co
F
Cu Ge
V
2 Se
Kr
Li Sr
B Ga Zr
Sc
1 Br Mo Sn Te Xe Ba Pb
As Rb Ru
Y Pd Cd Ce Pt
Nd Os
0 I Gd Dy Hg
Nb Sm Er Yb
Be Hf W Ir
Rh Ag Sb Cs La
−1 In Pr Au Tl Bi Th
Eu Ho U
Tb Re
Tm Lu
−2 Ta
0 10 20 30 40 50 60 70 80 90
Z (Element)
Abundances of Elements, Fig. 1 Solar system abundances (by number) of the 92 chemical elements, in a logarithmic
scale where log(N) = 6 for Silicon (from a compilation in Lodders 2003)
differences with the solar composition. The uni- See Also

versal predominance of H (90 % by number, but
70 % by mass) and He (9 % by number, but ▶ Nucleosynthesis, Neutrino
25 % by mass) and the relative abundances of ▶ Nucleosynthesis, Explosive
“metals” (to astronomers, elements heavier than ▶ Primordial Nucleosynthesis
He) is the most important similarity. On the other ▶ S-process
hand, the fraction of metals (metallicity, about ▶ Star
1.5 % in the Sun) appears to vary considerably
within the solar neighborhood (where the oldest
stars have a metallicity of 0.1 solar), across the
Milky Way disk (with young stars in the inner References and Further Reading
Galaxy having three times more metals than the
Sun), or in the galactic halo (with stellar Asplund M, Grevesse N, Sauval AJ, Scott P (2009) The
metallicities ranging from 0.1 to 0.00001 solar). chemical composition of the sun. Ann Rev Astron
Astrophys 47:481–522
These variations in composition reflect the his-
Lodders K (2003) Solar system abundances and conden-
tory of “chemical evolution” of the Milky Way sation temperatures of the elements. Astrophys
(Fig. 1). J 591:1220–1247
32 1989 AC
fabric present in metamorphic rocks and pro-

1989 AC duced by reorientation of minerals). Lens-shaped
boudins (cylinder-like structures making up a
▶ Toutatis layer in a deformed rock) of serpentinized ultra-
mafic rocks, up to several hundred meters long,
occur throughout the gneisses. No ca. 4.0–3.6 Ga
metasedimentary rocks have been discovered
Acasta Gneiss although sparse outcrops of locally tightly folded
quartzite, iron formation, and pelite are found
Samuel A. Bowring in the older gneisses. Weakly deformed, ca.
Department of Earth, Atmospheric and Planetary 3.6 Ga-old granitic dikes cut many outcrops.
Sciences, Massachusetts Institute of Technology, During the 1.88 Ga Calderian orogeny to the
Building 54-1126, Cambridge, MA, USA west, sheets of 1.9–2.5 Ga-old rocks were thrust
over the western edge of the Slave craton,
resulting in a set of north-trending folds and
Keywords metamorphism of underlying Archean rocks.
Ar-Ar biotite and U-Pb apatite dates record com-
Geochronology; Oldest rocks; Zircon plex reheating during this event at ca. 1.77 Ga.
The protoliths of the Acasta Gneiss range
from granite to tonalite/diorite in composition.
Definition Their U-Pb zircon dates indicate that the older
igneous crystallization ages are 4.03–3.96 Ga.
The Acasta Gneisses are the oldest known rocks Many zircons from all rock types contain older
on the surface of the Earth. They are exposed in cores with the oldest at 4.06 and 4.2 Ga, which is
northern Canada, north of Great Slave Lake, east consistent with the involvement of even older
of Great Bear Lake with the approximate position crust in their generation by partial melting or
of 65 100 N and 115 300 W. They have a com- assimilation. In general, the geochemistry of the
position close to granitic and are interpreted to Acasta Gneisses is not different from other
have formed, at least in part, from even older Archean and younger rocks: they are on average
rocks that may be as old as 4.2 Ga. enriched in light rare earth elements with variable
depletion in heavy rare earth elements, features
that are thought to reflect the presence of garnet in
Overview the source area. ▶ Radiogenic isotope systemat-
ics in whole rocks (Sm-Nd) and zircon (Lu-Hf)
The Acasta Gneisses are the oldest dated rocks on are also consistent with the involvement of older
Earth. They are exposed in northwestern Canada ▶ continental crust. Many of the rocks have zir-
(65 100 N and 115 300 W) along the western cons with thin overgrowths likely related to meta-
margin of the Archean Slave craton (>2.5 Ga), in morphism at ca 3.65 Ga, 3.6 Ga, and 3.4 Ga.
the core of a north-trending fold in the foreland of The formation and preservation of ▶ conti-
the Wopmay orogen, a 2.02–1.84 Ga-old oro- nental crust early in Earth’s history is of broad
genic belt. The Acasta Gneisses range in age interest to Earth scientists because the oldest con-
from 4.03 Ga to ca. 3.6 Ga with distinct groupings tinental crust provides a record of magma forma-
at 4.03–3.94 Ga, 3.74–3.72 Ga, and tion and the role of water in generating granitic
3.66–3.58 Ga. Rocks from these three distinct magmas over 4 billion years ago. The ca. 4 Ga
groups are compositionally diverse and range granitoids are very similar to those formed much
from ▶ granite to quartz diorite to tonalite. later in Earth’s history by plate-tectonic pro-
Rocks have been deformed several times cesses. No evidence of the late heavy bombard-
resulting in well-developed foliations (planar ment is preserved in the Acasta Gneisses.
Accretion Shock 33
See Also under a continental plate occurs, plate accretion is

the process of scraping oceanic floor sediments
▶ Canadian Precambrian Shield against the continental plate which form a prism A
▶ Continental Crust of material called accretionary wedge. Landmass
▶ Earth, Formation and Early Evolution accretion is the process of adding sediments
▶ Geochronology (alluvium) to a coastline or a riverbank, increas-
▶ Granite ing land area surface.
References and Further Reading See Also

Bowring SA, Housh TB (1995) The Earth’s early evolu-
▶ Magnetic Fields and Planetary Systems
tion. Science 269:1535–1540
Bowring SA, Williams IS (1999) Priscoan (4.00–4.03 Ga) Formation
orthogneisses from northwestern Canada. Contrib ▶ Planet Formation
Mineral Petrol 134:3–16 ▶ Plate Tectonics
Bowring SA, Housh TB, Isachsen CE (1990) The Acasta
gneisses: remnant of Earth’s early crust. Origin of the
earth. Oxford University Press, New York
Iizuka T, Horie K, Komiya T, Maruyama S, Hirata T,
Hidaka T, Windley BF (2006) 4.2 Ga zircon
xenocryst in an Acasta gneiss from northwestern Can- Accretion Shock
ada: evidence for early continental crust. Geology
34:245–248 Steven B. Charnley
Iizuka T, Komiya T, Ueno Y, Katayama I, Uehara Y,
Maruyama S, Hirata T, Johnson SP, Dunkley DJ
(2007) Geology and zircon geochronology of the 691, Astrochemistry Laboratory, NASA Goddard
Acasta Gneiss Complex, northwestern Canada: new Space Flight Center, Greenbelt, MD, USA
constraints on its tectonothermal history. Precambrian
Res 153:179–208
Definition
Generally, an accretion shock is a shock wave

occurring at the surface of a compact object or
Accretion
dense region that is accreting matter supersoni-
cally from its environment. In the context of
Daniele L. Pinti
astrobiology, an accretion shock is normally
GEOTOP Research Center for Geochemistry and
understood to mean the shock wave present at
Geodynamics, Université du Québec à Montréal,
the surface of the protosolar nebula or the
Montréal, QC, Canada
corresponding nebula surrounding a ▶ protostar,
as it accretes interstellar matter from the sur-
rounding molecular cloud.
Definition
In planetary sciences, accretion is the complex

process of formation of a planet, either rocky or See Also
gaseous, from the disk of dust and gas around a
protostar. In geology, accretion is a process by ▶ Protoplanetary Disk
which rocks and sediments are added to a tectonic ▶ Protosolar Nebula, Minimum Mass
plate (plate accretion) or a landmass (landmass ▶ Protostars
accretion). When subduction of an oceanic plate ▶ Shock, Interstellar
34 Accretion, Stellar
Definition
Accretion, Stellar
Acetaldehyde is an organic compound with
Steven W. Stahler the chemical formula CH3CHO
Department of Astronomy, University of
H O
California, Berkeley, CA, USA
H C C
Definition H H
It is the smallest aldehyde after formaldehyde. It
Stellar accretion refers to the inflow of ambient gas is a colorless liquid at room temperature with an
onto the surface of a star. During the process of star irritating odor. It can be obtained by the oxidation
formation, accretion builds up the object to its final of ▶ ethanol or by the reduction of ▶ acetic acid.
mass. The infalling gas is the interior portion of a When we drink alcohol, the ethanol is oxidized to
▶ dense core, a small ▶ molecular cloud that col- acetaldehyde, by alcohol dehydrogenase, which
lapses under the influence of its own gravity. The is then oxidized to acetic acid by aldehyde dehy-
object being built up in this manner is a protostar drogenase in the liver. It can be formed easily
and represents the first phase of stellar evolution. from gas mixtures containing methane by ultra-
Some infalling gas impacts the protostar directly. violet light and electric discharges, among others.
Much of the gas, however, has sufficient angular It reacts with hydrogen cyanide and ammonia to
momentum that it goes into orbit around the young give 2-aminopropionitrile, which gives ▶ alanine
star. The accreting gas thus creates a circumstellar (amino acid) after hydrolysis. It has been detected
disk. Matter spirals in through the disk onto the in extracts from carbonaceous chondrites. Melt-
surface of the protostar. The remaining part of the ing point: 123.5 C, boiling point: 20.2 C,
disk eventually gives rise to planets. density: 0.788 g cm 3.
See Also
See Also
▶ Dense Core
▶ Free-Fall Time ▶ Acetic Acid
▶ Gravitational Collapse, Stellar ▶ Alanine
▶ Molecular Cloud ▶ Aldehyde
▶ Protoplanetary Disk ▶ Chondrite
▶ Protostars ▶ Formaldehyde
▶ Protostellar Envelope
▶ Star Formation, Theory
Acetic Acid
Acetaldehyde
Kensei Kobayashi
Kensei Kobayashi Yokohama National University, Tokiwadai,
Yokohama National University, Tokiwadai, Hodogayaku, Yokohama, Japan
Hodogayaku, Yokohama, Japan
Synonyms Synonyms
Acetic aldehyde; Ethanal Ethanoic acid

Acetonitrile 35
Definition Definition
Acetic acid is a ▶ carboxylic acid with the chem- The organic compound acetone (CH3COCH3) is A
H O the simplest example of a ketone. Under standard
conditions, it is a colorless, flammable liquid.
ical formula CH3COOH H C C . Acetone is naturally produced by normal meta-
O H bolic processes in the human body. Since it is
H
miscible with water, it serves as an important
It is a colorless liquid at room temperature with
laboratory solvent. Rotational transitions in both
an irritating odor. Pure anhydrous acetic acid is
the ground vibrational state and in the first
sometimes called glacial acetic acid. It can be
excited torsional state have been detected by
obtained by oxidation of ▶ acetaldehyde, which
radio astronomers in ▶ molecular clouds.
occurs in human liver catalyzed by the enzyme
aldehyde dehydrogenase, or by the hydrolysis of
acetonitrile. It is easily formed in chemical evolu-
tion experiments, e.g., it was found among the
History
products of spark discharge experiment in a gas
Although detection of acetone in a molecular
mixture of methane, ammonia, hydrogen, and
cloud toward the center of our ▶ Milky Way gal-
water by S. L. Miller in 1953. It has been found in
axy was reported by radio astronomers in 1987,
extracts from carbonaceous ▶ chondrites and has
secure confirmation of its presence in interstellar
also been identified in ▶ molecular clouds. Melting
clouds was not achieved until some 15 years later.
point: 16.6 C, boiling point: 117.8 C, density:
1.0492 g cm3, acidity constant (pKa): 4.76.
See Also See Also
▶ Acetaldehyde ▶ Molecular Cloud

▶ Aldehyde ▶ Molecules in Space
▶ Carboxylic Acid ▶ Milky Way
▶ Chondrite
▶ Miller, Stanley References and Further Reading
▶ Molecular Cloud
Friedel DN, Snyder LE, Remijan AJ, Turner BE
(2005) Detection of acetone toward the orion-KL hot
core. Astrophys J 632:L95–L98
Acetic Aldehyde
▶ Acetaldehyde
Acetonitrile
Acetone
Kensei Kobayashi
William M. Irvine Yokohama National University, Tokiwadai,
University of Massachusetts, Amherst, MA, USA Hodogayaku, Yokohama, Japan
Synonyms Synonyms
Propanone CH3CN; Cyanomethane; Methyl cyanide

36 Acetylene
Acetonitrile is the simplest organic ▶ nitrile Acetylene is the simplest alkyne (hydrocarbons
with the chemical formula CH3CN that have a triple bond between two carbon
H atoms, with the formula CnH2n2). Under stan-
dard conditions in the laboratory, it is a colorless
C C N. It is a colorless liquid at but unstable gas. Because of its symmetry, linear
H of the form HCCH, it lacks a permanent electric
H dipole moment and hence has no allowed pure
room temperature with an ether-like odor. It can rotational transitions, making it undetectable at
be obtained by dehydration of acetamide or by millimeter wavelengths. Astronomers have
hydrogenation of a mixture of carbon monoxide observed its vibrational transitions in the infra-
and ammonia. It gives ▶ acetic acid and ammo- red, in both ▶ molecular clouds and in the enve-
nia after hydrolysis and gives ethylamine after lopes of evolved stars. It is an important link in
reduction. Acetonitrile itself is only slightly the chemistry of heavier carbon chain molecules
toxic but gives extremely toxic ▶ hydrogen cya- and related species in these regions. Acetylene is
nide by metabolism in the body. It is detected in also found as a minor component in the atmo-
▶ molecular clouds as an interstellar molecule and spheres of gas giants like the planet ▶ Jupiter, in
also found in cometary comas. When aminated on the atmosphere of Saturn’s satellite ▶ Titan, and
the methyl group, aminoacetonitrile is produced, in ▶ comets.
which is an important precursor of ▶ glycine. It is
completely miscible with water and often used as
an eluant in high-performance liquid chromatogra- History
phy (HPLC), with melting point, 45.7 C; boiling
point, 82 C; and density, 0.786 g cm3. Acetylene was discovered in 1836 by Edmund
Davy and then rediscovered in 1860 by French
chemist Marcellin Berthelot, who coined the
name “acetylene.” It was first observed in the
See Also
interstellar medium by Lacy et al. (1989) and in
▶ Comet Hyakutake and ▶ Comet Hale-Bopp
▶ Acetic Acid
(Brooke et al. 1996).
▶ Comet
▶ Glycine
▶ Hydrogen Cyanide
▶ Molecular Cloud See Also
▶ Nitrile
▶ Comet
▶ Comet Hale–Bopp
▶ Comet Hyakutake
▶ Molecular Cloud
Acetylene ▶ Molecules in Space
▶ Stellar Evolution
William M. Irvine
University of Massachusetts, Amherst, MA, USA
References and Further Reading
Brooke TY, Tokunaga AT, Weaver HA, Crovisier J,

Synonyms Bockelée-Morvan D, Crisp D (1996) Detection of
acetylene in the infrared spectrum of comet
Ethyne, HCCH Hyakutake. Nature 383:606–608
Acid Hydrolysis 37
Hartquist TW, Williams DA (1995) The chemically con-

trolled cosmos. Cambridge University Press, Achondrite
Cambridge
Lacy JH, Evans NJ II, Achtermann JM, Bruce DE, Arens A
JF, Carr JS (1989) Discovery of interstellar acetylene. Frank Sohl and Tilman Spohn
Astrophys J 342:L43–L46 Deutsches Zentrum f€ur Luft- und Raumfahrt (DLR),
Institut f€ur Planetenforschung, Berlin, Germany
Definition
Achiral
Achondrites are differentiated stony ▶ meteorites
Robert Hazen and constitute a minority among the stony meteor-
Geophysical Laboratory, Carnegie Institution of ites. The term literally means “without ▶ chon-
Washington, Washington, DC, USA drules” and therefore underlines the main
difference with ▶ chondrites. Achondrites are
igneous ▶ rocks or ▶ breccias of igneous
Synonyms rock fragments and thus their parent body has
experienced partial melting and recrystallization.
Mirror symmetric The class of achondrites includes the primitive
achondrites (e.g., ureilites) and achondrites in gen-
eral (e.g., aubrites, eucrites, howardites, diogenites,
Definition Martian meteorites, and lunar meteorites).
The term “achiral” is applied to any object – in

See Also
astrobiology most commonly a molecule, a
two-dimensional crystal surface, or a three-
▶ Breccia
dimensional crystal structure – that is invariant
▶ Chondrite
(i.e., superimposable) with its mirror image.
▶ Chondrule
Achiral objects possess a plane of symmetry,
▶ Meteorites
either a mirror or a glide plane symmetry
▶ Rock
operator. Common achiral objects include a
soccer ball, a pencil, and the letter “X,” in con-
trast with chiral objects such as a snail shell,
your left hand, and the letter “R.” Common
achiral molecules are H2O, CH4, and NH3 in
Acicular Ironstone
contrast with such chiral biomolecular species
▶ Goethite
as alanine and ribose. In chemistry, achiral should
not be confused with racemic, although in neither
case is the optical rotation of polarized light
affected. Acid Hydrolysis
Mark Dörr
See Also University of Southern Denmark, Odense M,
Denmark
▶ Chirality
▶ Enantiomeric Excess Definition
▶ Homochirality
▶ Racemic Mixture Hydrolysis (Greek: udor [hydor] = “water” and
▶ Stereoisomers lύsiB [lýsis] = “solution”) is a chemical reaction
38 Acid Maceration
in which a compound is cleaved by water. If a

proton-donating compound (Brønsted acid) Acidophile
catalyzes the reaction, it is called “acid hydroly-
sis.” Formally one part of the cleaved reaction Felipe Gomez
product receives a proton (H+), the other a Centro de Astrobiologı́a (CSIC/INTA),
hydroxyl (OH) moiety of a water molecule. Instituto Nacional de Técnica Aeroespacial,
Hydrolysis can also be catalyzed by a base. The Torrejón de Ardoz, Madrid, Spain
reverse reaction is called a “condensation
reaction.”
Keywords
Archaea; Chemolithoautotroph; Eukaryote; Iron

cycle; Prokaryote; Sulfur cycle
Acid Maceration
Emmanuelle J. Javaux
Definition
Palaeobiogeology-Palaeobotany-
Palaeopalynology, Geology Department,
Acidophiles are ▶ microorganisms that thrive
Université de Liège, Liège, Belgium
under acidic conditions, usually at very low pH
(<3). Natural niches where acidophiles can be
found are volcanic areas (Yellowstone), hydro-
Definition
thermal sources, deep-sea vents, metal mining
activities (Iron Mountain, Rı́o Tinto), or in the
Acid maceration is a technique used to extract
stomachs of animals. Acidophilic organisms can
organic-walled ▶ microfossils or kerogen from
be found in the different domains of the tree of
rock. A rock sample is cleaned to remove
life (▶ Archaea, Bacteria, and ▶ Eukarya).
external contamination and crushed into
small pieces. About 25 mg is macerated in
chlorhydric acid solution (HCl) to remove
Overview
carbonate minerals, rinsed with distilled water,
and then macerated in fluorhydric acid solution
The best characterized acidophiles belong mainly
(HF) to remove silicate minerals. A following step
to Bacteria and Archaea domains (Madigan and
of boiling the macerate in hot HCl removes fluo-
Martinko 2005). These microorganisms normally
rides formed during the previous acid step. This
are associated to sulfuric pools, acid mine drain-
protocol may vary according to the nature of the
age, or hydrothermal sources, that is, environ-
rock, of the fossils, and of their degree of preserva-
ments where sulfur compounds are present. The
tion. After neutralization of the final macerate with
origin of extreme acidic conditions is related with
distilled water, the residue is filtered on sieves of
the ▶ oxidation of reduced sulfur compounds
desired size fractions, then mounted on microscopic
which determines the strong relationship between
slides or kept in vials for other analyses.
acidophiles and sulfur chemistry. But also some
eukaryotes as the green algae Dunaliella
acidophila and the red algae Cyanidium
See Also caldarium (both organisms can live below pH
1.5) and some fungi are acidophiles. Surprisingly,
▶ Acritarch eukaryotic microbes are the principal contribu-
▶ Biomarkers, Morphological tors of biomass in some low-pH environments as
▶ Fossil ▶ Rio Tinto, a Spanish river which has a pH of
▶ Kerogen around 2 and contains much higher
Acidophile 39
concentrations of heavy metals than are typically cytoplasm at or near neutral pH (Kelch
found in fresh waters (López-Archilla et al. 2007). Therefore, intracellular proteins are
et al. 2001). Some interesting results from Rio not forced to develop acid stability through evolu- A
Tinto are related with evolutionary distances tion. However, some acidophiles, as Acetobacter
between acidophilic species that occur in the aceti, have an acidified cytoplasm. In the case of
river and their neutrophilic relatives. Authors acidophiles with acidic cytoplasm, all proteins are
reported some results that offer insight into adap- forced to evolve acid stability. Acetobacter has
tation to an extremely acidic environment. Adap- become as a good model for studying acid stability
tations associated with the transition from a mechanisms. Several authors have studied pro-
neutral to an acidic environment must occur rel- teins adapted to low pH and revealed that in
atively rapidly when measured on evolutionary acid-stable proteins there is an overabundance of
timescales (Amaral-Zettler et al. 2002). Authors acidic residues which minimizes low-pH destabi-
detected clones from the Rio Tinto that are lization induced by a buildup of positive charge.
closely related to neutrophilic species such as The relocation of acid-sensitive salt bridges to
Chlamydomonas noctigama, Chlorella regions with important functions in the unfolding
minutissima, and Colpidium campylum. From process is a very specialized case of acid stability
microscopic observations, rotifers were also for some proteins of acidophiles. But some others
identified as inhabitants of Rio Tinto, as well as mechanisms for protein stabilization in acid con-
heliozoans and other types of amoeba. ditions have been reported.
Acidophiles belonging to different orders of
the archaeal domain have been identified, such as
Sulfolobales (a particular order in the Applications
Crenarchaeota branch) and some facultative
anaerobic thermoacidophilic microorganisms as Acidophiles have industrial applications as in
Acidianus brierleyi and A. infernus and other biomining and bioremediation. Among the
thermophilic as Metallosphaera sedula, all of methods used for acid mine drainage treatment,
them related with metal mobilization in natural there are several which involve also metal-
environments and mining processes, immobilizing bacteria as a way for metal seques-
Thermoplasmatales (order included in tering. An important biotechnological application
Euryarchaeota branch), and some others groups of acidophiles is the process known as bioleaching
as nanoorganisms associated to Iron Mountain for metal extraction and exploitation of extremely
Mine and called ARMAN (Archaeal Richmond low-grade ores. Not only bacteria but also fungi
Mine Acidophilic Nanoorganisms). The have been used in related projects (Mohapatra
ARMAN group is composed by three different et al. 2007; Botuyan et al. 1996). Projects include
lineages deeply branched in the Euryarchaeota nickel extraction with A. ferrooxidans and Asper-
subgroup. gillus sp. fungi (Mohapatra et al. 2007) and sulfur
Acidophiles belonging to the bacterial domain removal from coal with Acidithiobacillus sp. or
are the phylum Acidobacteria, the order Aciditio- naturally present population bacteria on fossil
bacillales of ▶ Proteobacteria, the genus fuels as coal (Gómez et al. 1997).
Acidithiobacillus and Leptospirillum, and some
other related microorganisms as Acetobacter
aceti, a bacterium that belongs to the Acetobacter See Also
genus of Proteobacteria that produces acetic acid
(vinegar) from the oxidation of ethanol. ▶ Autotroph
Studies of mechanisms used for adaptation at ▶ Autotrophy
low pH by acidophiles reported interesting results ▶ Biodiversity
about efficient apparatus for pumping protons out ▶ Chemoautotroph
of the intracellular space as a way of maintaining ▶ Chemolithoautotroph
40 Acritarch
▶ Chemolithotroph
▶ Deep-Sea Microbiology Acritarch
▶ Early Archean
▶ Eukarya Emmanuelle J. Javaux
▶ Eukaryote Palaeobiogeology-Palaeobotany-
▶ Euryarchaeota Palaeopalynology, Geology Department,
▶ Extremophiles Université de Liège, Liège, Belgium
▶ Hot Spring Microbiology
▶ Hot Vent Microbiology
▶ Hydrothermal Environments Keywords
▶ Iron Cycle
▶ Microorganism Organic-walled microfossils
▶ Oxidation
▶ Prokaryote
▶ Proteobacteria Definition
▶ Pyrite
▶ Rio Tinto An acritarch is a microscopic organic-walled hol-
▶ Sulfur Cycle low vesicle with an unknown biological affinity.
▶ Yellowstone National Park, Natural Analogue The term comes from two Greek words: “acritos”
Site (unknown) and “arche” (origin). Acritarchs are
found in marine sediments from most of the geo-
References and Further Reading logical timescale, from the Archean to the pre-
sent, but were more abundant and diversified in
Amaral Zettler LA, Gómez F, Zettler E, Keenan BG, the Proterozoic and the Paleozoic. Their diversity
Amils R, Sogin ML (2002) Microbiology: eukaryotic decreased in the Mesozoic.
diversity in Spain’s River of Fire. Nature 417:137.
doi:10.1038/417137a
Botuyan MV, Toy-Palmer A, Chung J, Blake RC 2nd,
Beroza P, Case DA, Dyson HJ (1996) NMR solution Overview
structure of Cu(I) rusticyanin from Thiobacillus
ferrooxidans: structural basis for the extreme acid sta-
The term “acritarch” was coined by Evitt (1963)
bility and redox potential. J Mol Biol 263(5):752–767.
doi:10.1006/jmbi.1996.0613. PMID 8947573 to define any organic-walled ▶ microfossils that
Gómez F, Amils R, Marı́n I (1997) Microbial ecology cannot be assigned to a known biological group.
studies for the desulfurization of Spanish coals. Fuel The acritarchs form a polyphyletic artificial
Process Technol 52(1–3):183–189
group that may represent many different types
Kelch BA, Eagen KP, Erciyas FP, Humphris EL,
Thomason AR, Mitsuiki S, Agard DA (2007) Struc- of organisms, from bacteria to unicellular
tural and mechanistic exploration of acid resistance: (protists) or multicellular eukaryotes (e.g.,
kinetic stability facilitates evolution of extremophilic fungi, algae, animal eggs). Their size ranges
behavior. J Mol Biol 368(3):870–883. doi:10.1016/j.
from a few microns to a few millimeters,
jmb.2007.02.032. PMID 17382344
López-Archilla AI, Marin I, Amils R (2001) Microbial although most are microscopic. They are classed
community composition and ecology of an acidic into several artificial morphogroups based on the
aquatic environment: the Tinto River. Spain Microb morphology and presence and type of ornamen-
Ecol 41:20–35. doi:10.1007/s002480000044
tation of the vesicle. Filamentous organic-walled
Madigan M, Martinko J (eds) (2005) Brock biology of
microorganisms, 11th edn. Prentice Hall, Upper Sad- microfossils of unknown biological affinities are
dle River. ISBN 0-13-144329-1 not included among acritarchs. In the Paleozoic,
Mohapatra S, Bohidar S, Pradhan N, Kar RN, Sukla LB most acritarchs are assumed to represent cysts of
(2007) Microbial extraction of nickel from Sukinda
marine planktonic algae (phytoplankton).
chromite overburden by Acidithiobacillus
ferrooxidans and Aspergillus strains. Hydrometallurgy Acritarchs are very useful to date and correlate
85:1–8 marine sedimentary rocks (biostratigraphy),
Activated Nucleotide 41
▶ Fossilization, Process of
▶ Microfossils
▶ Proterozoic Eon A
Butterfield NJ, Knoll AH, Swett N (1994) Paleobiology of

the Neoproterozoic Svanbergfjellet formation Spits-
bergen. Fossils Strata 34:1–84
Evitt WR (1963) A discussion and proposals concerning
fossil dinoflagellates, hystrichospheres and acritarchs.
Proc Natl Acad Sci U S A 49:158–164
Grey K, Willman S (2009) Taphonomy of Ediacaran
acritarchs from Australia: significance for taxonomy
Acritarch, Fig. 1 Two acritarchs from the and biostratigraphy. Palaios 24:239–256
Mesoproterozoic (1.65 Ga Ruyang Group, China). Left, a Javaux EJ, Knoll AH, Walter MR (2001) Morphological
spiny acritarch (acanthomorph); right, a smooth-walled and ecological complexity in early eukaryotic ecosys-
acritarch (sphaeromorph) (Photograph: E Javaux) tems. Nature 412:66–69
Javaux EJ, Marshall CP, Bekker A (2010) Organic-walled
microfossils in 3.2-billion-year-old shallow-marine
especially in the late Proterozoic and the Paleo-
siliciclastic deposits. Nature 463:934–938
zoic, to reconstruct paleoenvironments, and to Knoll AH, Javaux EJ, Hewitt D, Cohen P (2006) Eukary-
assess the thermal maturity of the organic matter otic organisms in Proterozoic oceans. Phil Trans R Soc
(paleothermometer) and their potential as a source B 36:1023–1038
Servais T (1996) Some considerations on acritarch classi-
of hydrocarbon. Moreover, the detailed study of
fication. Rev Palaeobot Palynol 93:9–22
acritarch morphology, wall ultrastructure, and Strother PK (1996) Acritarchs. In: Jansonius J, McGregor
microchemistry gives precious information on the DC (eds) Palynology: principles and applications,
early evolution of eukaryotes in the Precambrian. vol 1. American Association of Stratigraphic Palynol-
ogists Foundation, Tulsa, pp 81–106
The oldest acritarchs known so far have
smooth walls (sphaeromorphs) and are preserved
in 3.2 Ga fine-grained detritic rocks deposited in
tidal shallow marine waters (Mesoarchean). The
oldest ornamented acritarchs (with concentric
striations) are 1.9 Ga and interpreted as probable
Acrylonitrile
eukaryotes. The oldest spiny acritarchs
▶ Vinyl Cyanide
(acanthomorphs) are unambiguously eukaryotic
and 1.65 Ga. Acritarchs can be preserved as flat-
tened vesicles in fine-grained detritic sediments
(shales, siltstones) or in three dimensions by
permineralization (partial or complete replace-
ment by minerals) in cherts or phosphorites. Activated Nucleotide
They are studied in thin sections or extracted
from the rock by ▶ acid maceration, depending Kunio Kawamura
on their state of preservation (Fig. 1). Department of Human Environmental Studies,
Hiroshima Shudo University, Hiroshima, Japan
See Also
Keywords
▶ Acid Maceration
▶ Eukaryotes, Appearance and Early Chemical evolution; Nucleotide; Oligonucleo-
Evolution of tide; RNA
42 Activated Nucleotide
Synonyms As mentioned above, the polymerization of

nucleotide 50 -triphosphates does not proceed in
Nucleoside 50 -monophosphorimidazolide the absence of an enzyme and a polynucleotide
template. Since enzymes are fairly complicated
biological catalysts, and since the first nucleic
Definition acids must have predated the first nucleic acid
templates, primitive pathways to form RNA
An activated nucleotide is a nucleoside polymers without an enzyme and a template mol-
50 -monophosphate possessing a leaving group, ecule have been investigated under simulated
such as imidazole, which provides sufficient energy Earth conditions. The activation of nucleoside
to form higher oligonucleotides. The activation of 50 -monophosphates is essential thermodynami-
nucleoside 50 -monophosphates by N-P bond forma- cally, since the nucleoside 50 -monophosphate
tion between the phosphate group and a base (such does not possess sufficient energy to form
as an amino acid or an imidazole) seems to be phosphodiester bonding. These monomers also
essential for the abiotic formation of RNA oligo- have the experimental benefit of reacting quite
mers. In contrast, DNA oligomers are less readily rapidly under laboratory conditions. Further-
formed from activated deoxynucleotides. more, the activation should have occurred from
plausible precursors under primitive Earth
conditions. Investigations by Orgel and
Overview coworkers have shown that the nucleoside
50 -monophosphate monomers, that is, nucleoside
The synthesis of ▶ RNA molecules may have 50 -monophosphoimidazolides and related com-
been an essential step for the emergence of life- pounds, can be considered as candidate primitive
like systems on the primitive Earth. In modern activated nucleotide monomers, as they can be
organisms, polymerization of RNA monomers formed under plausible primitive Earth condi-
only proceeds because thermodynamics and tions (Lohrmann and Orgel 1973; Orgel and
kinetics allow for biochemical polymerization Lohrmann 1974; Lohrmann 1977; Huang and
in organisms. In modern biology, nucleoside Ferris 2006). RNA oligomers up to 50 nucleotide
50 -triphosphate monomers are used to form units can be formed from the activated nucleotide
RNA polymers. This is thermodynamically monomers in the presence of a metal ion catalyst,
favorable, since nucleoside 50 -triphosphates pos- a clay mineral catalyst, or a polynucleotide tem-
sess high-energy phosphate groups. These by plate. In contrast, DNA oligomers do not form
themselves are rather kinetically slow to react in readily from the activated nucleotides.
solution; however, their polymerization becomes
kinetically favorable due to the involvement of
biological catalysts, RNA polymerases. In addi- See Also
tion, biological nucleotide polymerization is usu-
ally conducted on a DNA template; in the ▶ Nucleoside
absence of an RNA polymerase or a template, ▶ Nucleoside Phosphoimidazolide
the polymerization of 50 -triphosphate does not ▶ Oligonucleotide
proceed. ▶ RNA
RNA polymers are also formed in biology ▶ RNA World
from nucleoside 50 -diphosphate in the presence
of polynucleotide phosphorylase without a DNA
template. This is an exception, although this does
show that the polymerization of nucleoside Huang W, Ferris JP (2006) One-step, regioselective syn-
50 -diphosphates is also thermodynamically thesis of up to 50-mers of RNA oligomers by mont-
possible. morillonite catalysis. J Am Chem Soc 128:8914–8919
Active Asteroid 43
Lohrmann R (1977) Formation of nucleoside 50 -phosphor- where A is a constant called the frequency or
amidates under potentially prebiological conditions. pre-exponential factor, usually given in units of
J Mol Evol 10:137–154
Lohrmann R, Orgel LE (1973) Prebiotic activation s1, and R is the universal gas constant. A
processes. Nature 244:418–420
Orgel LE, Lohrmann R (1974) Prebiotic chemistry and
nucleic acid replication. Acc Chem Res 7:368–377 See Also
▶ Arrhenius Plot
Activation Energy
Henderson James (Jim) Cleaves II Active Asteroid

Earth–Life Science Institute (ELSI), Tokyo
Institute of Technology, Meguro–ku, Tokyo, Gerhard Hahn
Japan Asteroids and Comets, DLR, Institute of
Institute for Advanced Study, Princeton, NJ, Planetary Research, Berlin, Germany
USA
Blue Marble Space Institute of Science,
Washington, DC, USA Synonyms
Center for Chemical Evolution, Georgia Institute
of Technology, Atlanta, GA, USA Dual status objects; Mainbelt comet
Synonyms Definition
Ea Objects in typical asteroidal orbits, e.g., located

in the main asteroid belt, showing typical come-
tary activity, such as tails or comae, either tem-
Definition porarily or sporadically. The possible origin of
this “cometary” activity may either be intrinsi-
In chemical kinetics, the activation energy cally comet-like, e.g., outgassing, or triggered
(abbreviated Ea) is the energy barrier which externally, e.g., by impact events, collisions,
must be overcome for a sufficient number of etc. Such objects are not confined to the asteroid
reactant molecules to acquire enough kinetic belt, but exist also in the NEO population, and
energy for a reaction to occur appreciably. The among Centaurs. A comprehensive recent review
activation energy can generally be achieved by can be found in Jewitt (2012).
supplying energy, for example, in the form of
heat, to the system or through the intervention
of a catalyst. The Arrhenius equation gives the History
temperature dependence of the rate constant k: Ea
is obtained from the slope of a plot of ln k versus The first observation of such an object was the
1/T, where T is the absolute temperature in Kel- Centaur asteroid (2060) Chiron = 95P, which
vin. The activation energy is usually measured in was found in 1988–1989 to show unusual bright-
kJ/mol. ening that could be related to comet-like activity
Ea is related to the rate (k) via the equation exhibiting a coma. A member of the NEO
group that was observed as both an asteroid and
Ea ¼ RT lnðk=AÞ as a comet is (4015) Wilson-Harrington = 107P.
44 Active Site
Mainbelt asteroid (7968) Elst-Pizarro = 133P place. For example, in surface chemistry, this
shows repeated activity, most likely caused by could be a specific set of surface sites, and in
sublimation of water ice. biochemistry, it is a particular surface or cleft of
Most recent related finding is the detection of a protein ▶ enzyme or ▶ ribozyme surface where
water vapor on dwarf planet Ceres = asteroid catalysis occurs. In protein enzymes, the active
(1) Ceres. site is generally a pocket or cleft with specific
amino acid side chains presented in particular
orientations that bind a ▶ substrate and facilitate
See Also catalysis. Cofactors facilitating catalysis may
also be bound in the active site.
▶ Asteroid Belt, Main
▶ Centaurs (Asteroids)
▶ Comets, History of See Also
▶ Near-Earth Objects
▶ Catalyst
▶ Cofactor
References and Further Readings ▶ Enzyme
▶ Ribozyme
Jewitt D (2012) The active asteroids. Astron J 143:66 ▶ Substrate
(14 pp)
K€uppers M, O’Rourke L, Bockelee-Morvan D, Zakharov V,
Lee S, von Almen P, Carry B, Teyssier D, Marston A,
M€uller T, Crovisier J, Barucci MA, Moreno R (2014)
Localized sources of water vapour on the dwarf planet
(1) Ceres. Nature 505:525–527
List of Dual status object at the MPC: http://www.
minorplanetcenter.net/iau/lists/DualStatus.html. Last
Activity
accessed 5 May 2014
Henderson James (Jim) Cleaves II
Institute of Technology, Meguro–ku, Tokyo,
Japan
Active Site Institute for Advanced Study, Princeton, NJ,
USA
Henderson James (Jim) Cleaves II Blue Marble Space Institute of Science,
Earth–Life Science Institute (ELSI), Tokyo Washington, DC, USA
Institute of Technology, Meguro–ku, Tokyo, Center for Chemical Evolution, Georgia Institute
Japan of Technology, Atlanta, GA, USA
Institute for Advanced Study, Princeton, NJ,
USA
Blue Marble Space Institute of Science, Definition
Washington, DC, USA
Center for Chemical Evolution, Georgia Institute In chemistry, the activity of a species is a measure
of Technology, Atlanta, GA, USA of its effective concentration, as distinct from its
concentration. This is by convention usually
presented as a unitless value. Activity can be
Definition affected by factors such as pressure, temperature,
and the presence of other solutes. The difference
In catalysis, an active site is the specific location in value between concentration and activity is due
on the surface of a catalyst where reactions take to the fact that molecules in solution often display
Activity, Magnetic 45
nonideal behavior due to molecular interactions. History

In the gas phase, activity is called fugacity.
The Sun is the first known celestial X-ray source. A
It was discovered by rocket observations in 1949
See Also by a group of physicists in New Mexico, USA
(Friedmann et al. 1951). Since EUV photons and
▶ Oxygen Fugacity X-rays cannot penetrate the Earth’s atmosphere,
studies continued using high-altitude balloons
and finally satellites (14 launched in the world
between 1957 and 2006). The first solar X-ray
images were obtained on the Skylab space station
Activity, Magnetic
(1973–1979), establishing the link between
X-rays and ▶ magnetic fields. The Sun is now
Thierry Montmerle
continuously monitored by satellites on various
Institut d’Astrophysique de Paris, CNRS/
orbits, some of which are heliosynchronous, like
Université Paris 6, Paris, France
the recently launched NASA’s Solar Dynamics
Explorer. Following the progress in solar X-ray
imaging, the X-ray emission from individual
Keywords
stars was discovered by the Einstein satellite
(1978–1981), already showing “solar-type activ-
Chromosphere; Corona; Doppler-Zeeman imag-
ity” (flares), hence magnetic in origin. X-ray sat-
ing; Dynamo effect; Flares; Magnetic fields; UV
ellites currently in operation (NASA’s Chandra
radiation; X-rays
and ESA’s XMM-Newton) have detected hun-
dreds of thousands of stars with evidence for
magnetic activity.
Synonyms
Variability (magnetic)
Overview
Solar Activity
Definition The prototype of a “magnetically active star” is
the Sun, for which high-resolution images show,
Magnetic activity is a characteristic of low-mass at all wavelengths, numerous localized “active
stars, with the Sun as a prototype. Its most spec- regions” (the largest ones seen as sunspots in
tacular observational manifestation is in the form the visible domain). The solar magnetic activity
of stellar flares that can be seen in the optical, UV, is best seen at “hot” temperatures (105–107 K),
and X-ray domains. These flares correspond to i.e., in the EUV and X-ray domains. The
the release of magnetic energy itself temporarily corresponding regions of the Sun’s atmosphere
built up as a result of movements of the outer are, respectively, called the “chromosphere” and
layers of these stars, which are convective the “corona.” Magnetic activity is also seen in the
(“dynamo” effect). During flares, the stellar lumi- visible domain, which corresponds to the “pho-
nosity is enhanced by a factor which depends on tosphere” (the surface of the Sun as we see it with
the energy domain, but which is maximum in the naked eye). A closer view shows that the solar
X-rays (amplification by a factor of up to 10 or surface is constantly in motion: this is due to the
more). The whole phenomenon typically lasts a phenomenon of convection, which is the result of
few hours and may be described by a rapid energy transport from the interior of the Sun.
heating phase (minutes to tens of minutes), Figure 1 shows the optically visible Sun in its
followed by a long cooling phase (a few hours). ultraquiet epoch of Spring 2009, and Fig. 2
46 Activity, Magnetic
Activity, Magnetic, Fig. 1 The Sun in visible light,

Activity, Magnetic, Fig. 3 The Sun in X-rays, as seen
during its ultraquiet period of Spring 2009. Note the two
by the Japanese Hinode satellite. Active regions are again
minuscule black “spots”: they are the silhouettes of the
tightly linked with magnetic loops. Here the brightest
Space Shuttle servicing the International Space Station
regions reach a temperature of several 107 K (JAXA)
(Photograph NASA/T. Legaut)
shows for comparison the optical/EUV Sun,

which regained activity a few months later.
These structures evolve with time: on long-time
scales such as days, weeks (star spots, promi-
nences), but more spectacularly on timescales of
hours (flares, such as the one seen on the solar
limb of Fig. 2). Solar flares reveal material being
heated in a few tens of minutes and confined by
large magnetic loops; subsequently, the material
cools, decays, and eventually flows back to the
photosphere after a few hours. The magnetic field
is created via the so-called dynamo effect,
resulting from the macroscopic movement of
electrically charged convective gas. As shown
in Fig. 3, the contrast between “active” regions
and “quiet” regions is stronger at X-ray energies.
This turns out to be a crucial factor for studying
stellar magnetic activity.
Stellar Magnetic Activity

Activity, Magnetic, Fig. 2 The Sun in visible/EUV
light, having regained its activity in early 2010. The hot The so-called stellar magnetic activity is a char-
gas is confined by closed magnetic loops in many places, acteristic of all “solar-type stars” (by extension
in particular here at the upper left sector of the limb (color from the Sun: relatively cool stars with tempera-
coded: bright regions are the hottest, 106 K). There are
tures between 3,000 and 10,000 K and masses
also open magnetic field structures, which let the hot gas
escape in the form of the solar wind (Solar Dynamics between 0.1 M (1M = 1 solar mass) and
Observatory, NASA) 2 M; for comparison, the surface temperature
Activity, Magnetic 47
of the Sun is 6,000 K). These stars have con- A broader view of stellar magnetic activity is
vective outer layers and generate their own mag- provided by X-rays. In particular, the generally
netic field by the dynamo effect like the Sun. The large field of view of X-ray telescopes allows the A
crucial observational difference between these simultaneous study of all the stars present in the
stars and the Sun is that they cannot be resolved; field, especially if they belong to a cluster. This is
in other words, they appear as points in any imag- the case for young, pre-main sequence stars
ing device. However, one can reconstitute to (“▶ T Tauri” stars): as shown as an example in
some degree the stellar surfaces by using their Fig. 5, over 2,000 stars can be observed simulta-
rotation, which modulates the signal: since active neously in the Orion star formation region
regions (such as spots) are cooler, hence darker (associated with its famous nebula M42). Long
than the surrounding surface, their emission is duration observations (several days) reveal the
Doppler shifted when the spots appear to move occurrence of ubiquitous X-ray flares, most very
toward, then away from, the observer, as the star similar to solar flares, and a few being of much
rotates. In favorable cases, it is possible to go one longer duration and extreme temperatures (up to
step further, by using the Zeeman effect. This is a 108 K).
physical phenomenon in which, under the influ-
ence of magnetic fields, specific spectral lines are
slightly split into three components, sensitive to Basic Methodology
polarization, and which are also Doppler shifted
as a result of stellar rotation. Using sophisticated Solar and stellar magnetic activities are observed
deconvolution methods, one can then reconstruct primarily from space. The latest solar satellite,
not only the location and size of active regions NASA’s SDO (Solar Dynamics Observatory),
but also the topology of their associated magnetic launched in 11 February 2010, transmits high-
fields. Figure 4 shows such a reconstruction in the resolution full-disk optical/EUV images at the
case of a young solar-type star. unprecedented rate of 1 image per second
(Fig. 2). Meanwhile, ESA/NASA’s SOHO
(Solar and Heliospheric Observatory), launched
in December 1995, continues to provide data on
the Sun’s activity and solar wind in the same
domain, covering to this date more than one full
solar magnetic cycle of 11 years. The Sun is also
imaged in X-rays (Fig. 3), in particular by a series
of Japanese-led satellites (Yohkoh, now Hinode
since 2006), providing the best comparison set
for stellar magnetic activity studies. Stars are
observed using X-ray satellites (NASA’s
Chandra and ESA’s XMM-Newton, both
launched in 1999 and still in operation) that
have both a large field of view (a fraction of a
square degree) and high spatial resolution. In the
case of Chandra, the spatial resolution is compa-
rable with that of the best ground-based tele-
scopes (0.200 ), allowing astronomers to resolve
compact stellar clusters like those found in star-
forming regions. In addition to imaging, X-ray
Activity, Magnetic, Fig. 4 Image reconstruction of the
satellites also have spectral capabilities, allowing
active regions and magnetic field topology of the young
star SU Aur, observed via Zeeman-Doppler imaging measurement of densities, temperatures, and
(At Wikipedia: P. Petit, Observatoire Midi-Pyrénées) composition of the X-ray emitting plasma, and
48 Activity, Magnetic
Activity, Magnetic, Fig. 5 The Orion Nebula seen in correspondence holds between the stars visible on the
X-rays. Left: near-IR image of the Orion Nebula cluster, left and the point X-ray sources visible on the right.
which contains 2,000 stars (ESO/VLT/ISAAC). Right: A long exposure of this region shows ubiquitous flares
X-ray image of the same cluster, obtained with the from all the low-mass stars (From Montmerle et al. 2006)
Chandra satellite (NASA). An excellent, one-to-one
their dependence with time (for instance, flares phenomenon, which is the accretion of material
and associated coronal emission). from a circumstellar, protoplanetary disk, along
large-scale magnetic field lines connecting the
star and the disk. This phenomenon is seen as
Key Research Findings “soft” X-ray (<1 keV) emission due to a shock
when the accreted material falls onto the stellar
The vast majority of “solar-like stars” show evi- surface, superimposed on the “hard” (>1 keV)
dence for magnetic activity, and observations coronal emission.
show that X-rays are an excellent proxy to com-
pare with solar magnetic activity. However, the
level of activity is very different depending on Applications
stellar type. This level is generally measured by
the ratio of the energy output in X-rays (the X-ray For present-day human environment, the solar
luminosity LX) to the total energy output (the magnetic activity is strongly related to the notion
bolometric luminosity Lbol). For the Sun, LX / of “space weather,” i.e., the interaction of the
Lbol 107, with a variable amplitude of a factor solar wind with the Earth’s magnetic field. For
10 over magnetic cycles. In contrast, fully con- astrobiology, X-rays from the young Sun (and
vective stars, like “emission-line” stars, late-type from young stars in general) may have had an
cool stars, and very young T Tauri stars, have LX / important influence, via ionization of atoms and
Lbol reaching 103. This limiting factor is due to molecules, on early planetary atmospheres. More
“saturation,” i.e., when active regions generally, the study of solar and stellar magnetic
cover essentially all the stellar surface, as activity is related to fundamental plasma physics:
opposed to 5 % for the Sun. For the youngest low-density plasmas in the corona and (in the
▶ T Tauri stars (ages less than a few million case of young stars) interaction with a circum-
years), the magnetic activity is altered by another stellar disk and high-density plasmas in motion,
Adaptation 49
leading to the generation of magnetic fields in

outer convective zones (dynamo effect). Theoret- Adaptation
ical developments are now based on sophisticated A
numerical 3D simulations. Susanna Manrubia
Systems Biology Program, Centro Nacional de
Future Directions
From an astronomical point of view, the similar- Keywords

ities between solar and stellar magnetic activity
allow two different approaches to the same prob- Ecosystem; Environmental change; Genetic
lem, i.e., the origin of stellar magnetic fields: change; Mutation; Phenotype; Selection
(1) considering the “Sun as a star,” where prox-
imity allows detailed, localized studies, and
(2) considering “stars as Suns,” allowing statisti- Definition
cal approaches on magnetic activity properties as
a function of ▶ spectral type (mass, age, evolu- Adaptation is a dynamical process whereby
tionary status) for thousands of stars, in particular populations become better suited to their habitat. It
for young stars at the stage of planet formation. is promoted by ▶ environmental changes, be they
abiotic (e.g., climatic change) or biotic (e.g., the
appearance of a new trait in a predator or the extinc-
See Also tion of a competitor ▶ species). Adaptation is the
outcome of ▶ natural selection acting on heritable
▶ Faint Young Sun Paradox variation and leading to a change in the genetic
▶ Magnetic Field makeup of a population. It may involve changes in
▶ Sun (and Young Sun) any ▶ phenotypic trait, among others, in morphol-
▶ X-Rays (Stellar) ogy, physiology, dispersal, defense and attack mech-
anisms, development and growth, reproduction,
behavioral patterns, and ecological interactions.
Charles P, Seward FD (2010) Exploring the x-ray uni-

Overview
verse. Cambridge University Press, Cambridge
Feigelson ED, Montmerle T (1999) High-energy pro-
cesses in young stellar objects. Annu Rev Astron Adaptation has been an ongoing process ever
Astrophys 37:363 since the emergence of the first self-replicating
Friedmann H, Lichtman S, Byram E (1951) Photon coun-
molecules. Populations of replicating entities
ter measurements of solar x-rays and extreme ultravi-
olet light. Phys Rev 83:1025 generate continuous variability chiefly due to
G€
udel M (2007) The sun in time: activity and environ- ▶ mutations and (in the case of organisms) to
ment. Living Rev Sol Phys 4(3):1–137, http:// the migration of ▶ genes and other mobile geno-
solarphysics.livingreviews.org/Articles/lrsp-2007-3/
mic sequences. Adaptation is a gradual process
G€
udel M, Nazé Y (2009) X-ray spectroscopy of stars.
Astron Astrophys Rev 17:309 that occurs over many generations as the off-
Lanza AF (2009) Stellar coronal magnetic fields and star- spring (i.e., the genes) of individuals best suited
planet interaction. Astron Astrophys 505:339 to the current habitat become increasingly abun-
Montmerle T et al (2006) From suns to life: a chronolog-
dant. Under environmental changes, species can
ical approach to the history of life on earth 3 solar
system formation and early evolution: the first 100 mil- react in three different ways. In case of slow
lion years. Earth Moon Planet 98:39–95 change, they (1) perform habitat tracking if the
50 Adaptive Optics
change is exogenous (i.e., the population shifts ▶ Gene

with the ▶ environment to maintain the charac- ▶ Mutation
teristics of its habitat) or (2) undergo genetic ▶ Natural Selection
change. Only in the latter case are they truly ▶ Phenotype
adapting. If changes are too sudden, species can- ▶ Species
not adapt and (3) become extinct. Even if the
abiotic environment remains constant, the steady
generation of mutants within a population com- References and Further Reading
pels related species to change. The appearance of
teeth and claws in predators forces a simulta- Gould SJ (ed) (1993) The book of life: an illustrated
history of the evolution of life on Earth. WW Norton,
neous improvement in defense organs
New York
(as skeletons) to escape extinction. This sustained Maynard Smith J, Szathmáry E (1995) The major transi-
competition is called the Red Queen effect (Van tions in evolution. Oxford University Press, New York
Valen 1973; Stenseth and Maynard Smith 1984). Mayr E (1982) The growth of biological thought: diver-
sity, evolution and inheritance. Belknap Press of Har-
Many adaptive changes in the history of life are
vard University Press, Cambridge
gradual and involve a significant number of sequen- Moalem S, Prince J (2006) Survival of the sickest. Harper
tial modifications, as the evolution of tetrapod limbs Collins, New York
from the fins of precursor fish. Some changes (the Rose MR, Lauder GV (eds) (1996) Adaptation. Academic,
San Diego
invention of lungs, of vascular systems in terrestrial
Stenseth NC, Maynard Smith J (1984) Coevolution in
plants, or of organs for flight) permitted organisms ecosystems: red queen evolution or stasis? Evolution
to colonize new ecological niches (Gould 1993). 38:870–880
Certain major transitions in evolution (Maynard Van Valen L (1973) A new evolutionary law. Evol Ther
1:1–30
Smith and Szathmáry 1995) are probably not the
result of adaptation, but of contingency. Such might
be the case for the appearance of eukaryotic cells or
of multicellular organisms. Adaptive Optics
Since natural selection acts on the phenotype
as a whole, it is impossible to simultaneously Daniel Rouan
improve all its traits in the same degree (Mayr LESIA, Observatoire Paris-Site de Meudon,
1982). Also, traits that were once the result of Meudon, France
adaptation may turn to be disadvantageous when
the habitat changes. Hereditary hemochromatosis
causes an excessive absorption of iron that accumu- Definition
lates in human body tissues. This disease is more
common among Europeans: it protects against bac- Adaptive optics is a technique, used while
terial infections and probably became frequent at performing visible or infrared imaging from a
the time of the Black Death (Moalem and Prince ground-based telescope, that improves the
2006). This is an example of the many currently image quality that is otherwise degraded by the
nonadaptive traits that were permitted to thrive at atmospheric turbulence. The wave-front distor-
some point of time in the history of a species. tions, measured on the studied object or a nearby
star, are compensated by deforming a small, thin
mirror at a speed higher than turbulence (1 kHz
See Also typically) (Fig. 1). The deformable mirror is con-
jugated to the pupil of the telescope. The use of
▶ Colonization, Biological this technique is mandatory in any method, such
▶ Environment as coronagraphy, that aims at a direct imaging of
▶ Evolution, Biological exoplanets. Adaptive optics should not be con-
▶ Evolution, Molecular fused with active optics that corrects the large
Adenosine Triphosphate 51
Light from found in ▶ DNA and ▶ RNA. Via Watson-

telescope Crick base pairing in double-stranded ▶ DNA
and ▶ RNA, adenine forms two hydrogen bonds A
Adaptive
mirror
Distorted with ▶ thymine (T) and ▶ uracil (U), respec-
wavefront
tively. It is hydrolyzed to give ▶ hypoxanthine.
The half-life to hydrolysis in aqueous solution at
pH 7 is 1 year at 100 C and 6 105 years at
Beamsplitter 0 C. It has a UV absorption maximum at 260 nm.
Control Corrected
system wavefront
It has been found in the ▶ Murchison meteorite
and can be synthesized in HCN polymerizations,
▶ Fischer-Tropsch-type reaction, and electric
discharges acting on gas mixtures such as NH3-
CH4-C2H6-H2O.
High-resolution
Wavefront camera
sensor
See Also
Adaptive Optics, Fig. 1 Cartoon describing the princi-
ple of adaptive optics: the wave front deformed by atmo- ▶ DNA
spheric turbulence is corrected thanks to a deformable ▶ Fischer-Tropsch-Type Reaction
mirror that produces an inverse deformation several hun-
dred times per second, after a device called a wave-front ▶ HCN Polymer
sensor has measured the residual distortion. A real-time ▶ Hydrogen Cyanide
computer is used to analyze the residuals and to define the ▶ Hypoxanthine
proper surface to give to the mirror. The final improved ▶ Meteorite, Murchison
image is obtained on a camera
▶ Nucleic Acid Base
▶ RNA
primary telescope mirror deformed under grav-
▶ Thymine (T)
ity, and which is applied at a much lower fre-
▶ Uracil (Ura)
quency (0.1 Hz).
See Also
▶ Coronagraphy
Adenosine 50 -Triphosphatase
▶ Imaging
▶ ATPase
▶ Telescope
Adenine Adenosine Triphosphatase
Shin Miyakawa ▶ ATPase

Ribomic Inc., Minato-ku, Tokyo, Japan
Definition
Adenosine Triphosphate
Adenine (C5H5N5), with molecular weight
135.13, is one of the four nucleic acid bases ▶ ATP
52 Adiabatic Processes
Adiabatic Processes Adsorption
Lisa Kaltenegger William M. Irvine

Cornell University, Ithaca, NY, USA University of Massachusetts, Amherst, MA, USA
Definition
Definition
Adsorption is the accumulation of molecules
A process affecting a parcel of matter is said to from the gas phase, or more generally from any
be adiabatic if it occurs without addition or fluid (adsorption operates in liquids as well as
loss of heat from the parcel. No exchange of gases), onto a solid surface, e.g., of an ▶ inter-
heat means that temperature changes in the stellar dust grain. The molecules may be bound
parcel must be due to changes in pressure by physical (van der Waals), electrostatic, or
alone. In planetary atmospheres, the adiabatic chemical (e.g., hydrogen bonding) forces.
lapse rate is the change in air temperature
with changing height, resulting from
pressure change. The so-called dry adiabatic See Also
lapse rate has the slope dðln TÞ=dðln pÞ ¼ R=cp ,
where T is temperature, p is pressure, R is the ▶ Interstellar Dust
specific gas constant (which depends on the mean
molecular weight of the mixture), and cp is the
specific heat at constant pressure. For
Earth’s atmosphere, the value R/cp corresponds
to about 2/7. Adiabatic processes are called AEB
isentropic, i.e., they leave entropy unchanged,
provided that the changes in state of the Michel Viso
system are slow enough that the system CNES/DSP/SME, Vétérinaire/DVM, Astro/
remains close to thermodynamic equilibrium at Exobiology, Paris Cedex 1, France
all times.
Synonyms
See Also Agência Espacial Brasileira; Brazilian space

agency
▶ Atmosphere, Model 1D
▶ Atmosphere, Structure
▶ Atmosphere, Temperature Inversion Definition
▶ Exoplanets, Modeling Giant Planets’
Atmospheres Numerous committees since 1961 have been
▶ Grey Gas Model charged with space activities in Brazil. In Febru-
▶ Non-Grey Gas Model: Real Gas ary 1994, the Brazilian Space Agency was
Atmospheres established within the Department of Science
Aerobic Mesophilic Bacterial Spores 53
and Technology. The agency is partnering with See Also

four institutes in charge of specific missions: the
National Institute for Space Research (INPE) in ▶ Aerobic Respiration A
charge of developing satellites and products for ▶ DNA Damage
civilian use, the Institute of Aeronautics and ▶ Electron Acceptor
Space in charge of developing planes and the ▶ Respiration
launch vehicles, and two launching bases in
Alcantara and Barriera do Inferno. The space
agency is developing numerous cooperation
with Europe, Japan, China and the United States.
Aerobic Mesophilic Bacterial Spores
For further information: http://www.aeb.gov.br/
Catharine A. Conley
NASA Headquarters, Washington, DC, USA
Aerobe
Definition
Felipe Gomez
Centro de Astrobiologı́a (CSIC/INTA), Instituto “Spores” or more precisely “bacterial endo-
Nacional de Técnica Aeroespacial, Torrejón de spores” are resistant dormant bodies produced
Ardoz, Madrid, Spain by some ▶ microorganisms (Gram-positive bac-
teria) upon exposure to stressful environmental
conditions. Aerobic microorganisms grow while
exposed to oxygen, while mesophilic microor-
Definition
ganisms grow on nutrient-rich media at tempera-
tures comfortable for humans (roughly between
Aerobes are organisms that can tolerate or require
15 C and 40 C). Spores are capable of surviving
the presence of (strict aerobe in this case) oxygen.
extreme environmental conditions in a dormant
Oxygen is an extremely strong oxidant and
state and proliferating when introduced into a
produces very reactive radicals, which react
hospitable environment. This particular type of
with amino acids or nucleic acids inactivating the
spores is commonly used in ▶ planetary protec-
functional sites of enzymes or producing
tion as reference microorganisms for the qualifi-
lethal mutations. Practically, all animals are
cation of ▶ bioburden reduction processes.
aerobes; most fungi and many prokaryotes
Numbering the spores per unit of surface is also
can survive in the presence of oxygen. To do so,
used as a proxy when measuring the relative clean-
aerobic organisms require the presence of
liness of spacecraft components and systems.
detoxification activities, like catalases and peroxi-
dases. Among aerobes there are different kinds of
organisms: obligate aerobes, which require oxy- See Also
gen for growth and use oxygen as final ▶ electron
acceptor in the ▶ respiration process; facultative ▶ Aerobe
aerobes, which can use oxygen or not, to obtain ▶ Bacteria
energy; microaerophiles, which require low levels ▶ Bioburden
of oxygen; and aerotolerants, which are not ▶ Spore
affected by the presence of oxygen. ▶ Survival
54 Aerobic Respiration
Overview
Aerobic Respiration
The atmosphere presents a series of challenges
Juli Peretó for life, from radiation to ▶ desiccation. The
Institut Cavanilles de Biodiversitat i Biologia absolute amount of solar radiation and the pro-
Evolutiva, Universitat de València, València, portional contribution of ultraviolet-B and
Spain ultraviolet-C radiation increase with altitude,
both of which are particularly hazardous to bio-
molecules. The low temperature and pressure at
Synonym 29 km above the surface of the Earth are similar
to those on ▶ Mars and create problems due to
Oxygen respiration freezing and desiccation. Finally, the lack of
nutrient availability in the atmosphere creates an
additional challenge for life.
Definition The ▶ survival of airborne microbes should
not be confused with growth and division while
Aerobic ▶ respiration is a respiration in which airborne. In fact, one of the critical questions that
dioxygen (O2) serves as the terminal ▶ electron has yet to be answered unequivocally is, “do
acceptor of an electron transport chain. microbes metabolize and divide while airborne?”
If they do, then the atmosphere may be consid-
ered a true habitat rather than just a place where
See Also they are transient interlopers.
Given the hostility of the environment, Earth’s
▶ Anaerobic Respiration atmosphere just above the surface contains a
▶ Electron Acceptor variety of airborne microorganisms that are
▶ Respiration thought to originate from the soil, lakes, oceans,
animals, plants, as well as any process causing
▶ aerosols or dust. The numbers of viable air-
borne microbes recovered from the atmosphere
Aerobiology vary seasonally with the highest numbers
obtained during the summer and fall and the
Rocco Mancinelli lowest in the winter. The distances that airborne
Bay Area Environmental Research Institute, organisms may travel range from a few kilome-
NASA Ames Research Institute, NASA Ames ters to thousands of kilometers. There is a statis-
Research Center, Moffett Field, CA, USA tically significant positive correlation between
the total number of viable ▶ bacteria isolated
from urban air and the concentration of
Keywords suspended particulate matter in the air. The
organisms may be protected from drying by
Biology, of the atmosphere; Microorganisms, in adsorbed water on the surfaces of these
the atmosphere; Pollen; Spores suspended particles.
Studies of the biology of the upper tropo-
sphere and lower stratosphere (5–20 km) date
Definition back to the late 1800s using balloons. But these
studies are few in number and not well controlled.
Aerobiology is the study of the occurrence, The organisms collected included fungi and
movement, and dispersal of living or once-living ▶ spore-forming bacteria. Later studies reported
material through the atmosphere. a larger variety of nonspore-forming microbes,
Aerosols 55
especially a variety of pigmented bacteria. How- Mandrioli P, Caneva G, Sabbioni C (eds) (2003) Cultural
ever, the composition and prevalence of microor- heritage and aerobiology – methods and measurement
techniques for biodeterioration monitoring. Kluwer,
ganisms in the middle to upper troposphere the Netherlands, 245 pp A
(8–15 km altitude) and their role in aerosol-
cloud precipitation interactions are unresolved.
Using meteorological rockets, fungi and
pigmented bacteria have been isolated from as Aeronautics and Space Agency of
high as 77 km, the highest altitude from which FFG
microbes have been isolated. These studies, how-
ever, all used culturing methods to determine ▶ ASA
microbial counts. It has been estimated that
those methods allow studying only between
0.1 % and 10 % of the total microbial biota in
any given environment. Therefore, it is specu-
lated that a number of microbes may exist in the Aerosols
upper atmosphere that we do not have the ability
to culture and go unnoticed and uncounted. More François Raulin
recently, however, studies using molecular Faculté des Sciences et Technologie, Université
microbiology techniques, microscopy, and con- Paris Est Créteil et Paris Diderot, LISA – UMR
ventional culturing have shown that bacterial CNRS 7583, Creteil, France
cells outnumber fungal cells in the upper
troposphere.
Synonyms
See Also Atmospheric dusts; Atmospheric particles; Haze

particles
▶ Aerosols
▶ Bacteria
▶ Desiccation Definition
▶ Extreme Environment
▶ Mars Aerosols are small liquid or solid particles in
▶ Solar UV Radiation, Biological Effects suspension in a gas. Solid smoke particles from
▶ Spore the burning of vegetation, dust formed by wind
▶ Survival erosion of soil, or liquid droplets produced by the
ocean waves are examples of aerosols.
Aerosols are present in many planetary envi-
ronments, for example, in the atmospheres of
Mars, the giant planets, and ▶ Titan, and they
DeLeon-Rodriguez N, Latham TL, Rodriguea LM, were probably present in the primitive atmo-
Barasexh JM, Anderson BE, Beyersdorf AJ, Ziemba sphere of the Earth, much as they are presently.
LD, Bergon M, Nenes A, Konstantinidis KT Atmospheric aerosols can play an important role
(2013) Microbiome of the upper troposphere: species
in climate, producing an antigreenhouse effect,
composition and prevalence, effects of tropical storms,
and atmospheric implications. Proc Natl Acad Sci U S like the haze in Titan’s atmosphere. Atmospheric
A 110:2575–2580 photochemistry in several extraterrestrial envi-
Horneck G, Klaus D, Mancinelli RL (2010) Space micro- ronments, like Titan, produces organic aerosols
biology. Mol Microbiol Rev 74:121–156
which are similar to laboratory-synthesized
Lacey ME, West JS (2007) The air spora – a manual for
catching and identifying airborne biological particles. ▶ tholins, which can produce a complex set of
XV. Springer, Dordrecht prebiotic chemicals when reacted with water.
56 Affinity Chromatography
See Also See Also
▶ Atmosphere, Structure ▶ Antibody

▶ Carbon ▶ Chromatography
▶ Cassini ▶ Enzyme
▶ Cassini-Huygens Space Mission ▶ HPLC
▶ Clouds ▶ Substrate
▶ Flux, Radiative
▶ Hydrocarbons
▶ Jupiter
▶ Methane Cuatrecasas P, Wilchek M, Anfinsen CB (1968) Selective
▶ Refractory Molecule enzyme purification by affinity chromatography. Proc
▶ Saturn Natl Acad Sci U S A 61:636–643. doi:10.1073/
▶ Tholins pnas.61.2.636
▶ Titan
Affinity Constant
Affinity Chromatography
Mark Dörr Earth–Life Science Institute (ELSI), Tokyo
University of Southern Denmark, Odense M, Institute of Technology, Meguro–ku, Tokyo,
Denmark Japan
Institute for Advanced Study, Princeton,
NJ, USA
Definition Blue Marble Space Institute of Science,
Washington, DC, USA
Affinity ▶ chromatography is a (bio-) chemical Center for Chemical Evolution, Georgia Institute
separation method based on highly specific of Technology, Atlanta, GA, USA
molecular interactions (affinity) such as between
antigens and antibodies, ▶ enzymes and ▶ sub-
strates, or receptors and ligands. The stationary Synonyms
phase is commonly composed of beads of a gel
(e.g., agarose gel) with a covalently bound ligand Association constant; Binding constant
(e.g., an ▶ antibody). Affinity chromatography is
one of the most powerful separation methods, as
it combines the size fractionation capability of Definition
gel permeation chromatography with specific,
reversible interactions of molecules. In chemistry and biochemistry, the affinity con-
stant is the reciprocal of the dissociation constant,
where both are equilibrium constants describing
History the strength of binding between a catalyst such as
an ▶ enzyme or ▶ ribozyme and its ▶ substrate.
The method was introduced by Cuatrecasas, A Km value is a specific example of an affinity
Wilchek, and Anfinsen in 1968 (Cuatrecasas constant in enzymatic reactions. For example, the
et al. 1968). Pedro Cuatrecasas and Meir Wilchek equilibrium for the formation of an enzyme-
were jointly rewarded the Wolf Prize in Medicine substrate (ES) complex between an enzyme (E)
1987 for this discovery. and a substrate (S),
Akilia 57
Ex þ S y $ Ex S y ; (1)
Agência Espacial Brasileira
can be represented as A
▶ AEB
jEx Sy j
Km ¼ (2)
½Ex ½Sy
€ r Luft- und Raumfahrt der
Agentur fu
where [E], [S], and [ES] are the concentrations of FFG
enzyme, substrate, and the enzyme-substrate
complex, respectively, and x and y represent ▶ ASA
their stoichiometric coefficients.
The affinity constant, also known as the
Michaelis constant, has units of per molar
(M1) or l/mole. Affinity constants can vary sig- Agenzia Spaziale Italiana
nificantly with solution conditions (e.g., temper-
ature, pH, and ionic strength). ▶ ASI
This equilibrium is also the ratio of the rate of
association (kass) and rate of dissociation (kdiss).
Two different enzyme-substrate complexes may AIB
have the same affinity constants, but one could
have a high kass and kdiss, while the other may ▶ Aminoisobutyric Acid
have a low kass and kdiss.
See Also Akilia
▶ Enzyme Minik T. Rosing

▶ Ribozyme Nordic Center for Earth’s Evolution, Natural
▶ Substrate History Museum of Denmark, University of
Copenhagen, Copenhagen, Denmark
Definition
AFGL 915, IRAS 06176-1036
Akilia is the name of a small island (~2.3 km by
▶ Red Rectangle 1.4 km) south of the town of Nuuk on the SW
coast of ▶ Greenland (63.933 N, 51.667 W).
The Akilia sequence of rocks has its name from
this island, but they occur throughout the
AGB Eoarchean of West Greenland. The Akilia
sequence consists mainly of granitic gneiss and
▶ Asymptotic Giant Branch Star includes some of the oldest rocks on Earth. The
largest member of this group of rocks is the
ca. 3.8 Ga-old ▶ Isua supracrustal belt composed
of metamorphosed pillow basalts, clastic, and
Age Measurement chemical sediments that constitute the oldest
known supracrustal sequence. Fractionated C,
▶ Geochronology N, and S isotopic compositions of materials of
58 Al-Andalus, Cosmological Ideas
probable sedimentary origin are thought to be movement, it violated significant principles of

biogenic and to provide the oldest record of life Aristotelian physics. Objecting to the Ptolemaic
on Earth. model required the production of different geo-
metrical solutions that would allow for singular
movement around the Earth and would explain
the observations. Ibn Tufayl claimed to have
See Also
arrived at a solution and promised to write it in
a separate book but apparently never had the time
▶ Earth, Formation and Early Evolution
to do so. Yet, Ibn Tufayl’s student, al-Bitruji,
▶ Greenland
credited his master by providing the basis for
▶ Isua Supracrustal Belt
the former’s solution.
Ibn Tufayl’s objections had their roots in the
works of Ibn Sīnā, who inspired much of Ibn
Tufayl’s works, including his most famous
Al-Andalus, Cosmological Ideas novella “Philosophus Autodidactus.” Ibn Sīnā
also rejected Ptolemy’s model and claimed to
Ahmed Ragab have arrived at a solution, but after much effort
Harvard Divinity School, Cambridge, MA, USA he decided not to share it with any of his students
and not to write it. Ibn Sīnā’s student, al-Juzjānī,
composed a solution for this problem in a letter
Keywords addressed to his master. However, Juzjānī’s solu-
tion was not sufficient.
Islam; Islamic astronomy; Andalus; Iberia; Medi- Ibn Tufayl’s more prominent student, Ibn
eval; Ibn Tufayl; Ibn Rush; Averroes; al-Bitruji; Rushd (Averroes; d. 1198), wrote an epitome to
Arzachel; al-Zarqālī; Ibn Sīnā Ptolemy’s Almagest and acquired much of his
fame because of his detailed commentaries on
Aristotle. In his commentaries on Aristotle’s
Overview metaphysics and De Caelo, Ibn Rushd criticized
the Ptolemaic model and considered it to be
Ibn Tufayl’s (Abubacer; d. 1185) main engage- unnatural. He explained that Aristotle and the
ment with planetary theories was centered on his ancients had arrived at perfect solutions for the
rejection of Ptolemaic eccentrics and epicycles as celestial movement that escaped Ptolemy and
they contradicted the basic principles of Aristo- that these solutions should be rediscovered.
telian physics. Ptolemy’s eccentrics and epicy- While he claimed to have worked on these solu-
cles implied two main violations of the classical tions in his youth, Ibn Rushd left the task unfin-
principles of natural philosophy. On one hand, ished to his successors.
the eccentric meant that planets moved around a Ibn Tufayl’s other student, al-Bitruji (d. 1204),
center different from the Earth. On the other who seemed to have worked entirely indepen-
hand, epicycles meant that planets moved west dently from Ibn Rushd without either of the two
to east in their epicycles while moving east to being aware of the other, was a more proficient
west around the eccentric, therefore introducing astronomer and produced a model in which he
double movement, which contradicted the single attempted to address his master’s objections.
simple movement explained in works of natural Bitrūjī maintained a homocentric universe in cor-
philosophy. While Ptolemy’s model came close respondence with natural philosophy, eliminating
to solving the observational variation in eccentrics and epicycles; he developed the notion
Alanine 59
of rotating poles that was mentioned by Aristotle Saliba G (2002) Greek astronomy and the medieval arabic
in De Caelo, even wondering why Ptolemy did tradition the medieval islamic astronomers were not
merely translators. They may also have played a key
not propose such a solution. Both Ibn Rushd and role in the copernican revolution. Am Sci 360–367 A
al-Bitrūjī accepted this polar movement, which
was developed by Theon of Alexandria into spiral
movement.
In addition to such cosmological discussions,
the Toledan Tables, predicting the positions of
Alanine
the Sun, Moon, and planets, represented a more
important contribution of Andalusian astrono-
Jeffrey Bada
mers in fields of practical and computational
Scripps Institution of Oceanography, La Jolla,
astronomy. About a dozen astronomers in
CA, USA
Toledo, the most prominent of whom was the
astronomer and instrument-maker al-Zarqālī
(Arzachel; d. 1087), and which included both
Definition
Muslims and Jews, undertook different mathe-
matical and astronomic calculations that resulted
Alanine is one of the 20 a-amino acids found in
most prominently in the tables. Al-Zarqālī had an
coded proteins with the formula C3H7NO2 and
important theoretical contribution in relation to
H
trepidation (a medieval theory related to the pre- O H
cession of the equinoxes), and he also improved the structure C C N .
Ptolemy’s calculations of the length of the HO H
Mediterranean. CH3
The a-carbon of alanine is chiral so there are
two optical isomers (▶ enantiomers), l- and
d-alanine. Alanine appears to be one of the first
and is the second most common amino acids in
Carmody FJ (1952) The planetary theory of Ibn Rushd. terrestrial proteins. It was the first ▶ amino acid
Osiris 10:556–586 synthesized in the laboratory, by Adolph Strecker
Goldstein RB (1972) Theory and observation in medieval in 1850, who reacted acetaldehyde with hydrogen
astronomy. Isis 63(1):39–47
cyanide and ammonia in aqueous solution. It is
Ibn Tufayl, Abu Bakr Muhammad, Abu’l Walid Muham-
mad Ibn Rushd (1999) Two Andalusian Philosophers. also readily produced in ▶ spark-discharge
Translated by Jim Colville. Kegan Paul International, experiments from a reduced gas mixture of meth-
New York ane, ammonia, and hydrogen and has been found
Langermann Y T (1997) Arabic cosmology. Early Sci
in carbonaceous chondrites. Alanine is known to
Med 2(2):185–213
Nasr SH, Leaman O (1996) History of Islamic Philosophy, stabilize helical structures in proteins.
Routledge History of World Philosophies. Routledge,
London
Sabra AI (1998) Configuring the universe: aporetic, prob-
lem solving, and kinematic modeling as themes of
See Also
arabic astronomy. Perspect Sci 6(3):288–330
Sabra AI (1984) The Andalusian revolt against ptolemaic ▶ Amino Acid
astronomy: averroes and Al-Bitruji. Transform Tradit ▶ Chirality
Sci 133–53
▶ Enantiomers
Saliba G (1994) Early arabic critique of ptolemaic cos-
mology: a ninth-century text on the motion of celestial ▶ Spark Discharge
spheres. J Hist Astron 25:115 ▶ Strecker Synthesis
60 Albedo
4pR2 sT 4eq ¼ ð1 AÞpR2 F; (1)

Albedo
which equates the energy thermally radiated by
Mark S. Marley the planet (left side) to the stellar (solar) energy
NASA Ames Research Center, Moffett Field, absorbed. The Bond albedo is frequently (as in
CA, USA Eq. 1) computed by integrating the total reflec-
tivity of a planet over the incident flux from the
parent star and thus depends both on the reflec-
Keywords tivity spectrum and the spectral type of the star.
A planet that efficiently scatters in the blue and
Clouds; Equilibrium temperature; Planet; absorbs in the red portion of the spectrum will
Spectroscopy thus have a higher integrated Bond albedo when
illuminated by a blue star than by a red star, since
a greater proportion of the incident light is
Synonyms scattered away in the former case. However, the
Bond albedo can be defined as a function of
Reflectivity wavelength, A(l), e.g., Irvine et al. (1968), and
is then usually termed the spherical albedo.
The geometric albedo is defined as the ratio of a
Definition planet’s reflectivity measured at zero phase angle
(opposition) to that of a Lambert disk (which has
Albedo is a unitless measure of the reflectivity of the same apparent brightness at all viewing angles)
an object. Albedo can range between zero and of the same radius. The geometric albedo is a
one. Several different types of albedos have been function of wavelength and, because it is measured
defined, and it is important to appreciate their at opposition (when the phase angle f = 0), does
unique characteristics. not require information on the dependence of scat-
tering with phase. For a perfectly reflecting Lam-
bert sphere, the geometric albedo is two-thirds and
for a semi-infinite purely ▶ Rayleigh scattering
History atmosphere it is three-fourths. Both such idealized,
perfectly scattering objects would have a Bond
Albedo is derived from the Latin “albus” or albedo of 1, but the latter atmosphere sends more
“white.” The term was first applied to optics light directly back to the observer at zero phase
by Johann Heinrich Lambert (who gave his angle and thus has a higher geometric albedo. The
name to the Lambert disk) in his text Photometria geometric albedo is a fixed quantity for a given
in 1760. planet, so the computation of the geometric albedo
does not depend on the type of incident flux. Other
types of albedos have been defined. Great care
Overview thus must always be exercised to be certain that
the correct albedo is being discussed.
From a planet-wide perspective, the albedo of
most importance is the Bond albedo, A, the ratio
of incident energy reflected into all angles by a
planet to the total incident energy received from See Also
its star. The Bond albedo appears in the equation
for the equilibrium temperature of a rapidly rotat- ▶ Atmosphere, Structure
ing planet Teq, with radius R receiving an incident ▶ Clouds
flux F, ▶ Rayleigh Scattering
al-Bı̄rūnı̄, Abū Rayḥān 61

al-Bı̄rūnı̄, Abū Rayḥān
Cahoy K, Marley M, Fortney J (2010) Exoplanet albedo
spectra and colors as a function of planet phase, sepa- A
Ahmed Ragab1 and Allyssa Metzger2
ration, and metallicity. Astrophys J 724:189–214 1
de Pater I, Lissauer J (2010) Planetary sciences, 2nd edn. Harvard Divinity School, Cambridge, MA, USA
2
Cambridge University Press, Cambridge Harvard University, Cambridge, MA, USA
Hanel RA, Conrath BJ, Jennings DE, Samuelson RE (1992)
Exploration of the Solar System by Infrared Remote
Sensing. Cambridge University Press, Cambridge
Irvine WM, Simon T, Menzel DH, Pikoos C, Young AT Keywords
(1968) Multicolor photometric photometry of the
brighter planets, III. Astron J 73:807–828 Islam; Islamic astronomy; Avicenna; Medieval;
Seager S (2010) Exoplanet atmospheres. Princeton Uni-
India; Zij; Astrology; Calendar; Chronology
versity Press, Princeton
Overview
Albedo Feature
Abū Rayḥān al-Bīrūnī (d. 1048) was a Persian
Daniela Tirsch scholar of mathematics, astronomy, astrology,
German Aerospace Center DLR, Institute of and geography, among other disciplines.
Planetary Research, Berlin, Germany Although he changed patrons frequently, he pro-
duced his most important works in the Ghaznavid
court.
Synonym Educated in the dominant Greco-Islamic
scholarship, Bīrūnī was interested in comparing
Regio different scientific traditions. His “Chronology of
Ancient Nations” was a detailed exposition of
calendars and chronologies used by different
Definition nations, including Arabs, Greeks, Persians,
Jews, and Syriacs, among others. The Chronol-
A geographic area on the surface of a ▶ planet or ogy was unprecedented in its comparative
satellite that is distinguished from adjacent ter- approach and its careful examination of different
rains by a difference in brightness or ▶ albedo. chronological systems. It also included detailed
Classical albedo features have been identified descriptions of these nations’ different religious
with Earth-based telescopes, while no detailed mor- feasts.
phology could be resolved. Time-varying albedo While accompanying the Ghaznavid sultan
features can be due to seasonal changes, for exam- Maḥmūd (r. 998–1002) in India, Bīrūnī came in
ple, due to frost covers or due to the redistribution of contact with Indian scholars and composed a
bright dust or dark sand sheets by aeolian activity. significant volume where he addressed Indian
Thanks to high-resolution imagery on board of space religious beliefs and rituals, Indian sciences
probes, albedo feature nomenclatures are increas- including astronomy and cosmology, and Indian
ingly replaced with detailed feature descriptions. chronology and calendars, which were missing
from his Chronology. Bīrūnī’s work was the first
to discuss “foreign” nations and histories in
See Also non-polemical fashion with the stated goal of
understanding different scientific and belief tra-
▶ Albedo ditions. His work in India included transmitting
▶ Planet Ptolemy’s Almagest and other books of Greco-
▶ Regio Islamic sciences to Indian scholars. Bīrūnī’s
62 Alcohol
comparative view was based on his stated belief References and Further Reading
that sciences are universal and they speak to
universal laws of nature, only in different lan- Bag AK (1975) Al-Biruni on Indian arithmetic. Indian
J Hist Sci 10(2):174–184
guages and using different models. Comparison
Berggren JL (1985) The origins of Al‐Bīrūnī‘S “Method of
of scientific theories, and of calendars and chro- the Zijes” in the theory of sundials. Centaurus 28(1):1–16
nologies, was meant to explain the different Hartner W, Mathias S (1963) Al-Biruni and the theory of
approaches to a single universal truth. the solar apogee (an example of originality in Arabic
science). In: Scientific change. London: Heinemann,
Bīrūnī’s most influential text in the Islamic
p 206–218
context was his al-Qānūn al-Masʾūdī (The Langermann YT (1997) Arabic cosmology. Early Sci Med
Mas‘udic Canon), an 11-volume book in which 2:185–213
he developed algebraic solutions of third-degree Pines S (1964) The semantic distinction between the terms
astronomy and astrology according to Al-Biruni. Isis
equations and distinguished between the motions
55(3):343–349
of precession and the solar apogee. While the Rosenthal F (1976) Al-Biruni between Greece and India.
Canon was primarily based on Ptolemy’s Alma- Paper presented at the Biruni symposium
gest, it was also an attempt at incorporating ele- Saliba G (1994) Early arabic critique of ptolemaic cos-
mology: a ninth-century text on the motion of celestial
ments of Indian and Persian astronomy in the
spheres. J Hist Astron 25:115
dominant Greco-Islamic tradition. Bīrūnī was Saliba G (1995) A history of arabic astronomy: planetary
aware of some difficulties in the Ptolemaic plan- theories during the golden age of Islam. NYU Press,
etary theory. However, contrary to his contempo- New York
Saliba G (2002) Greek astronomy and the medieval arabic
raries, Bīrūnī’s objections did not only stem from
tradition the medieval Islamic astronomers were not
Ptolemaic violations of the principles of classical merely translators. They may also have played a key
natural philosophy, as Avicenna’s (d. 1037) role in the copernican revolution. Am Sci 90:360–367
objections were, but were based on contradic-
tions between the Ptolemaic system and Bīrūnī’s
own observations. Bīrūnī’s correspondence with
Avicenna showed some of Bīrūnī’s skepticism Alcohol
and doubts on Aristotelian natural philosophy.
While working as a court astrologer, Bīrūnī Kensei Kobayashi
composed his Tafhī m on astrology, which he Yokohama National University, Tokiwadai,
started with lengthy discussions of geometry Hodogayaku, Yokohama, Japan
and astronomy, with one quarter of the book for
the actual discussion of judicial astrology. He
argued that a well-trained astrologer, worthy of Definition
the name, must be proficient in all these
sciences. As part of his systematic observations, An alcohol is an organic compound containing a
Bīrūnī was interested in astronomical hydroxyl (-OH) group. In the IUPAC nomencla-
instruments – he wrote five books on instruments, ture system, the suffix “-ol” is used in the name of
including a lengthy one on the astrolabe. He also each alcohol. The simplest alcohol is ▶ methanol
described a coordinated viewing of the (methyl alcohol, CH3OH), and the most com-
lunar eclipse on 24 May 997 with Abu al-Wafa’ monly encountered alcohol is ▶ ethanol (ethyl
al-Buzjani (d. 998 CE) – from Khwarazm alcohol, C2H5OH) which is contained in alco-
and from Baghdad – to better calculate holic beverages. Alcohols with three or more
coordinates of major Islamic cities. Some of carbons have isomers: Propyl alcohol has two
Bīrūnī’s minor works discuss astronomical- isomers, which are propan-1-ol
mathematical concepts such as chords (Istikhrāj (CH3CH2CH2OH) and propan-2-ol (CH3CH
al-awtār), “shadows” or tangents (Ifrād (OH)CH2). Many low-molecular-weight alco-
al-maqāl), and planetary transits (Tamhīd hols, such as ethanol and propan-2-ol, are used
al-mustaqarr). as disinfectants, since they diffuse easily through
Aldose 63
cell membranes and denaturize proteins. Com- See Also

pounds whose hydroxyl group is connected to
benzene ring are referred to as phenols. ▶ Acetaldehyde A
▶ Aldose
▶ Amino Acid Precursors
See Also ▶ Carboxylic Acid
▶ Formaldehyde
▶ Ethanol ▶ Formose Reaction
▶ Methanol ▶ Glyceraldehyde
▶ Propanal
▶ Propionaldehyde
Aldehyde
John H. Chalmers
Scripps Institute of Oceanography Geosciences
Research Division, University of California, San Aldose
Diego, La Jolla, CA, USA
Definition Institute of Technology, Meguro–ku, Tokyo,
Japan
Aldehydes are organic compounds containing the Institute for Advanced Study, Princeton, NJ,
RCHO functional group where R can be hydro- USA
gen or another carbon-containing moiety. Alde- Blue Marble Space Institute of Science,
hydes are named after the corresponding Washington, DC, USA
carboxylic acids by dropping the -ic or -oic suffix Center for Chemical Evolution, Georgia Institute
and adding -aldehyde or -al (e.g., acetic acid of Technology, Atlanta, GA, USA
▶ acetaldehyde, ethanoic acid ! ethanal), or if
derived from acyclic aliphatic hydrocarbons, by
dropping the final e and adding -al (e.g., propane
! propanal). Formyl derivatives of ring com- Definition
pounds may be called carbaldehydes as in
cyclopentanecarbaldehyde. See the IUPAC rules An aldose is a ▶ monosaccharide where one end
for more complex cases. of the molecule is sp2 hybridized and thus pre-
Aldehydes can be reduced to alcohols or oxi- sents as a CHO or ▶ aldehyde group. These are
dized to carboxylic acids and undergo a variety of typically reactive ends of the molecules
addition and condensation reactions, for example, which may be involved in redox or addition
addition of cyanide in the cyanohydrin synthesis, or reactions. Examples of biologically important
condensation with another aldehyde in a benzoin aldoses include ▶ ribose and glucose.
reaction or condensation with the carbon adjacent ▶ Glycolaldehyde is the simplest aldose. One of
to an aldehyde or ketone in the aldol reaction. the most important aldoses is ribose, which
plays key roles as part of the pentose
phosphate cycle in the form of ribose 5-phosphate
History as well as the backbone sugar element of
ribonucleic acid (RNA). Another aldose deriva-
The name may be derived from the phrase tive of ribose, 2-deoxyribose, plays the same role
“▶ alcohol dehydrogenatum.” in DNA.
64 Algae
See Also along with cyanobacteria dominate global oce-

anic primary production.
▶ Aldehyde
▶ Carbohydrate
▶ Glycolaldehyde History
▶ Monosaccharide
▶ Ribose The term “blue-green algae” is a colloquial term
and refers to cyanobacteria that are members of
the domain Bacteria. This term is seldom used in
current literature but is still frequently encoun-
tered in popular articles and other media.
Algae
Linda Amaral-Zettler
Overview
Marine Biological Laboratory, Josephine Bay
Paul Center for Comparative Molecular Biology
Algae are differentiated by various means,
and Evolution, Woods Hole, MA, USA
including phylogenetic affiliation, photosynthetic
pigment type, and morphological features includ-
ing: the number of flagella, external ornamenta-
Keywords
tion (e.g., scales, frustules), subcellular
ultrastructure, and colony-forming abilities.
Brown algae; Coccolithophorids; Diatoms; Dino-
They are polyphyletic and possess representa-
flagellates; Euglenoids; Golden algae; Green
tives in several major lineages of the Eukarya
algae; Phytoplankton; Red algae; Seaweeds
including alveolates (e.g., dinoflagellates),
chlorarachniophytes (e.g., Chlorarachnion),
cryptomonads, euglenids, glaucophytes (e.g.,
Synonyms
Cyanophora), haptophytes (e.g., Emiliania
huxleyi), red algae (Rhodophyta), stramenopiles
Photosynthetic eukaryotes; Protists with
(e.g., diatoms, brown algae), and Viridiplantae
chloroplasts
(e.g., Chlorophyta).
Newer taxonomic classifications based on
molecular phylogenetic methods employing
Definition multigene approaches place representatives
from these lineages into higher taxonomic level
The algae are an eclectic grouping of photosyn- groupings called supergroups (Keeling
thetic eukaryotes that ranges from microscopic et al. 2005). These six major supergroups are
picoeukaryotes (1 mm) (Courties et al. 1994) to the “Plantae,” “Chromalveolata,” “Rhizaria,”
macroscopic multicellular seaweeds (50 m) (Sze “Excavata,” “Amoebozoa,” and
2003). Algae are phylogenetically and morpho- “Opisthokonta” – the first four of these contain
logically diverse, occur in benthic and planktonic algal representatives. Plantae (also referred to as
forms, and can be free living, symbiotic, preda- the Archaeplastida (Adl et al. 2005)) include the
tory, or parasitic. They inhabit diverse environ- red algae, green algae, and streptophytes;
ments including several extreme environments of Chromalveolata include alveolates and
astrobiological interest, such as desert varnish, stramenopiles; Excavata include euglenids;
permafrost, and highly acidic Mars analog envi- Rhizaria include chlorarachniophytes that group
ronments (Seckbach 2007). Green algae have within the Cercozoa. A note of caution is that the
unicellular members that share common ancestry resilience of these supergroups has been called
with land plants (Charales). Eukaryotic algae into question (Parfrey et al. 2006; Yoon
ALH 84001 65
et al. 2008), so it is important to take this into Parfrey LW et al (2006) Evaluating support for the current
consideration when using these terms. Of these classification of eukaryotic diversity. PLoS Genet
2(12):e220
six major supergroups, only the Opisthokonta Seckbach J (ed) (2007) Algae and cyanobacteria in A
appear to be strongly supported in robust phylo- extreme environments. Springer, Dordrecht, p 811
genetic analyses. Sze P (2003) A biology of the algae, 4th edn. McGraw-
In addition to chlorophyll a, algae are further Hill, Boston
Yoon HS et al (2008) Broadly sampled multigene trees of
distinguished on the basis of other types of pho- eukaryotes. BMC Evol Biol 8(1):14
tosynthetic pigments they possess. Alveolates,
cryptomonads, haptophytes, and stramenopiles
also contain chlorophyll c, while chlorarach- Internet Resources
niophytes, euglenids, Viridiplantae, and The Tree of Life Project: http://tolweb.org/
cryptomonads contain chlorophyll b. Other AlgaeBase: http://www.algaebase.org/
accessory pigments, such as phycobilins, further
distinguish cryptomonads, glaucophytes, and red
algae.
A feature that all algae share is the ability to
photosynthesize. There is strong evidence that ALH 84001
this characteristic was the result of a single endo-
symbiotic event that occurred between a cyano- Jean-Pierre de Vera
bacterium and an ancestor of the glaucophytes, DLR, Institut f€ur Planetenforschung, Berlin,
red algae, and green algae (including plants) Germany
(Keeling 2010). The uptake of both green and
red algae by other eukaryotes has occurred mul-
tiple times in what is referred to as “secondary Synonyms
symbioses.” In some dinoflagellates, tertiary
symbioses can occur via plastid replacement. Allan Hills 84001
Plastids are also sometimes stolen and used by
their host for brief periods of time by sea slugs,
dinoflagellates, as well as other protists, such as
ciliates and foraminifera – this process is called Definition
kleptoplasty.
ALH 84001 (abbreviation of Allan Hills 84001) is
a 1.93 kg ▶ meteorite found in 1984 on the Allan
See Also Hills ice field, Antarctica (Victoria Land), by US
meteorite searchers. ALH 84001 has been classi-
▶ Eukarya fied as ▶ achondrite and is thought to be from
▶ Mars. It mainly consists of coarse-grained
cataclastic orthpyroxene-rich material and
References and Further Reading among the ▶ SNC meteorites defines the class
of SNC-orthopyroxenites. In 1996, NASA scien-
Adl SM et al (2005) The new higher level classification of tists announced that the meteorite might contain
eukaryotes with emphasis on the taxonomy of protists.
▶ fossils of Martian microorganisms, a view that
J Eukaryot Microbiol 52(5):399–451
Courties C et al (1994) Smallest eukaryotic organism. has been widely criticized. Radiometric dating
Nature 370(6487):255–255 suggests that ALH 84001 is 4.1 billion years
Keeling PJ (2010) The endosymbiont origin, diversifica- old. The piece of ▶ rock has been ejected from
tion and fate of plastids. Phil Trans R Soc
Mars by an impact event 15 million years ago.
B 365:729–748
Keeling PJ et al (2005) The tree of eukaryotes. Trends Thirteen thousand years ago, the meteorite
Ecol Evol 20(12):670–676 landed on Earth.
66 Alignment of Dust Grains
See Also
Aliphatic Carboxylic Acids
▶ Achondrite
▶ Bacteria ▶ Fatty Acids, Geological Record of
▶ Fossil
▶ Mars
▶ Meteorites
▶ Rock
▶ SNC Meteorites Aliphatic Hydrocarbon

Alignment of Dust Grains Institute of Technology, Meguro–ku, Tokyo,
Japan
William M. Irvine Institute for Advanced Study, Princeton, NJ,
University of Massachusetts, Amherst, MA, USA USA
Washington, DC, USA
Definition
▶ Interstellar dust produces not only extinction of
transmitted starlight but also introduces polariza-
tion of that light, with a positive correlation Definition
between the amount of reddening and the linear
polarization. This effect is normally ascribed to An aliphatic hydrocarbon is an organic com-
the alignment of asymmetric grains in the galactic pound composed of carbon and hydrogen which
magnetic field. When the direction of alignment does not contain aromatic rings. It may be linear
changes along the line of sight, a circularly polar- or cyclic and may contain unsaturated double or
ized component is produced. Consequently, obser- triple bonds; thus, alkanes, alkenes, and alkynes
vations of this polarization provide (model- are all aliphatic compounds. Some illustrative
dependent) information on both dust grain proper- examples are ▶ methane, ethylene, ▶ acetylene,
ties and on the galactic magnetic field. Various and cyclopentane.
mechanisms have been proposed to produce the
grain alignment. Since circularly polarized light
could conceivably affect the chiral symmetry of
irradiated molecules such as amino acids, it could See Also
possibly play a role in producing the observed
▶ enantiomeric excess in some meteoritic organics, ▶ Acetylene
although this is far from being demonstrated. ▶ Aromatic Hydrocarbon
▶ Methane
See Also
▶ Chirality
▶ Enantiomeric Excess Alkaline Lakes
▶ Interstellar Dust
▶ Reddening, Interstellar ▶ Soda Lakes
Alkaliphile 67
Alkaliphiles require alkaline environments

Alkaliphile and, in most cases, sodium ions for their growth,
germination, and sporulation (Kudo and A
Antonio Ventosa and Rafael R. de la Haba Horikoshi 1983). Isolation of alkaliphilic micro-
Department of Microbiology and Parasitology, organisms in laboratory conditions must be car-
Faculty of Pharmacy, University of Sevilla, ried out in alkaline media containing sodium
Sevilla, Spain carbonate, sodium bicarbonate, or sodium
hydroxide, following conventional means.
Alkaliphiles are widely distributed in different
Keywords habitats and isolated from soils, feces, and
alkaline and/or saline lakes. The frequency of
Alkaline; Extreme habitat; Extremophile; pH; alkaliphilic microorganisms in neutral “ordinary”
Soda lake soil samples is 102–105/g of soil, which
corresponds to 1/10 to 1/100 of the population
of the neutrophilic microorganisms (Horikoshi
Definition 1991). Some studies show that alkaliphilic
bacteria have also been found in deep-sea
Alkaliphiles are microorganisms that grow opti- sediments collected from depths of up to
mally or very well at pH values above 9, often 10,898 m in the Mariana Trench
between 10 and 12, but cannot grow or grow (Takami et al. 1997).
slowly at the near-neutral pH value of 6.5 Most alkaliphiles have an optimal growth at
(Horikoshi 1999). around pH 10, which is the most significant dif-
ference from well-investigated neutrophilic
microorganisms. These alkaliphilic microorgan-
Overview isms can grow in such extreme environments
because their internal pH is maintained at
There is no precise definition of what character- 7.5–8.5, despite a high external pH of 8–13
izes an alkaliphilic organism. Several microor- (Aono et al. 1997). Therefore, one of the key
ganisms exhibit more than one optimum pH for features in alkaliphily is associated with the cell
growth depending on growth conditions, particu- surface, which discriminates and maintains the
larly nutrients, metal ions, and temperature. intracellular neutral environment separate from
However, the definition given above is the most the extracellular alkaline environment.
extended one. Alkaliphiles have two mechanisms of cytoplas-
Many different taxa are represented among the mic pH regulation. The first one involves the cell
alkaliphiles, including ▶ prokaryotes (aerobic wall structure, which contains acidic polymers
▶ bacteria belonging to the genera Bacillus, that function as a negatively charged matrix and
Micrococcus, Pseudomonas, and Streptomyces; may reduce the pH value at the cell surface (Aono
▶ anaerobic bacteria from the genera and Horikoshi 1983). The surface of the cytoplas-
Amphibacillus, Anaerobranca, and Clostridium; mic membrane must presumably be kept below
halophilic ▶ archaea belonging to the genera pH 9, because the cytoplasmic membrane is very
Halorubrum, Natrialba, Natronomonas, and unstable at alkaline pH values (pH 8.5–9.0)
Natronorubrum; methanogenic archaea from the much below the pH optimum for growth (Aono
genus Methanohalophilus; anaerobic archaea et al. 1992). The second strategy to maintain
from the genus Thermococcus; cyanobacteria; pH ▶ homeostasis consists of the use of the
spirochetes; actinomycetes; sulfur-oxidizing and Na+/H+ membrane antiporter system (Dc
sulfate-reducing bacteria), eukaryotes (▶ yeasts dependent and DpH dependent), the K+/H+
and filamentous ▶ fungi), and even phages antiporter, and ATPase-driven H+ expulsion
(Horikoshi 1998, 1999). (Krulwich et al. 1998).
68 Alkanoic Acids
The flagella motility of alkaliphiles is consid- Horikoshi K (1999) Alkaliphiles: some applications of
ered to be driven by a sodium-motive force their products for biotechnology. Microbiol Mol Biol
Rev 63:735–750
instead of a proton-motive force, as shown by Kobayashi T, Uchimura K, Miyazaki M, Nogi Y,
neutrophiles. These alkaliphiles are most motile Horikoshi K (2009) A new high-alkaline alginate
at pH 9.0–10.5, whereas no motility is observed lyase from a deep-sea bacterium Agarivorans
at pH 8; in addition, they require Na+ for motility sp. Extremophiles 13:121–129
Krulwich TA, Ito M, Hicks DB, Gilmour R, Guffanti AA
(Horikoshi 1998). (1998) pH Homeostasis and ATP synthesis: studies of
Studies of alkaliphiles have led to the discov- two processes that necessitate inward proton translo-
ery of many types of enzymes that exhibit inter- cation in extremely alkaliphilic Bacillus species.
esting properties. Alkaliphilic microorganisms Extremophiles 2:217–222
Kudo T, Horikoshi K (1983) Effect of pH and sodium ion
produce some enzymes such as proteases, amy- on germination of alkalophilic Bacillus species. Agric
lases, cyclomaltodextrin glucanotransferases, Biol Chem 47:665–669
pullulanases, cellulases, lipases, xylanases, Takami H, Inoue A, Fuji F, Horikoshi K (1997) Microbial
pectinases, chitinases, and alginate lyases that flora in the deepest sea mud of the Mariana Trench.
FEMS Microbiol Lett 152:279–285
are of great interest (Horikoshi 1999; Kobayashi
et al. 2009).
See Also Alkanoic Acids
▶ Anaerobe ▶ Fatty Acids, Geological Record of

▶ Archaea
▶ Bacteria
▶ Cyanobacteria
▶ Eukaryote Allan Hills 84001
▶ Fungi
▶ Homeostasis ▶ ALH 84001
▶ Methanogens
▶ Prokaryote
▶ Soda Lakes
▶ Yeast ALMA
Thijs de Grauuw
ALMA, Vitacura, Santiago, Chile
Aono R, Horikoshi K (1983) Chemical composition of cell
walls of alkalophilic strains of Bacillus. J Gen
Microbiol 129:1083–1087 Synonyms
Aono R, Ito M, Horikoshi K (1992) Instability of
the protoplast membrane of facultative
alkaliphilic Bacillus sp. C-125 at alkaline pH values Atacama large millimeter array; Atacama large
below the pH optimum for growth. Biochem millimeter/submillimeter array
J 285:99–103
Aono R, Ito M, Horikoshi K (1997) Measurement of
cytoplasmic pH of the alkaliphile Bacillus lentus
C-125 with a fluorescent pH probe. Microbiology Definition
143:2531–2536
Horikoshi K (1991) Microorganisms in alkaline environ- The Atacama Large Millimeter/submillimeter
ments. Kodansha-VCH, Tokyo
Array (ALMA) is an international radio telescope
Horikoshi K (1998) Alkaliphiles. In: Horikoshi K, Grant
WD (eds) Extremophiles: microbial life in extreme on a dry site at 5,000 m elevation in the Atacama
environments. Wiley-Liss, New York, pp 155–179 Desert of northern Chile. The US$1.4 billion
Alpha Helix 69
ALMA project is a partnership of Europe, Japan, of 0.518. The component A of this system is a
and North America in cooperation with the Repub- G2V star (see Spectral Type) with a mass of 1.1
lic of Chile. ALMA is funded in Europe by the solar-masses, a luminosity of 1.519 times solar, A
European Organization for Astronomical and an effective temperature of 5,790 K. Its com-
Research in the Southern Hemisphere (ESO), in ponent B has a spectral type of K1V and its mass,
Japan by the National Institutes of Natural Sci- luminosity, and effective temperature are equal to
ences (NINS) in cooperation with the Academia 0.934 solar-mass, 0.5 solar luminosity, and
Sinica in Taiwan, and in North America by the 5,214 K, respectively.
National Science Foundation in cooperation with In 2012, a team of scientists lead by Xavier
the National Research Council of Canada and the Dumusque announced the detection of a 1.13
National Science Council of Taiwan. Construction Earth-mass planet around a Cen B. One year
and operation of the facility are led on behalf of later, the existence of this planet, known as a
Europe by ESO, on behalf of Japan by the National Cen Bb, was questioned in an article by Artie
Astronomical Observatory of Japan (NAOJ), and Hatzes (2013).
on behalf of North America by the US National The probable existence of a Cen Bb would
Radio Astronomy Observatory (NRAO), which is indicate that, unlike the region around a Cen
managed by the Associated Universities, Inc. A where terrestrial planet formation encounters
(AUI). The telescope consists of 66 high-precision complications, as predicted by several researchers,
antennas operating over the instrument wave- planet formation around a Cen B may be efficient.
length range of 0.3–10 mm, with 25 provided by
the North American partners, 25 by European col-
laborators, and 16 by the Asian collaborators. All References and Further Reading
of the antennas can work together as a single
telescope. ALMA is fully operational since the Dumusque X, Pepe F, Lovis C et al (2012) Nature 491:207
Guedes JM, Rivera EJ, Davis E et al (2008) Astrophys
end of 2013.
J 679:1582
Hatzes AP (2013) Astrophys J 770:133
Th’ebault P, Marzari F, Scholl H (2009) MNRAS 393:L21
Alpha Centauri Bb
Nader Haghighipour Alpha Helix

Institute for Astronomy, University of Hawaii-
Manoa, Honolulu, Hawaii, HI, USA John H. Chalmers
Scripps Institute of Oceanography Geosciences
Research Division, University of California,
Definition San Diego, La Jolla, CA, USA
Alpha Centauri Bb is a putative planet orbiting

the star Alpha Centauri B. Alpha Centauri is a Definition
triple stellar system consisting of a moderately
close binary a Cen AB, and a distant M dwarf The alpha (or a-) helix is one of the two most
companion known as Proxima Centauri at common ▶ polypeptide secondary structural
approximately 15,000 AU away from the binary. motifs and consists of a right-handed helix with
At a distance of 4.37 light years from the Sun, a 3.6 amino acid residues per turn. The helix has a
Centauri is the closest stellar system to the solar pitch of 0.54 nm and a width of 1.2 nm and is
system and is located in the southern constella- stabilized by ▶ hydrogen bonds between the pep-
tion of Centaurus. The binary system has a tide -C=O- and -NH- moieties of every fourth
semimajor axis of 23.5 AU and an eccentricity bond. The amino acids proline and glycine tend to
70 Alpha Particles
kink or break alpha helices, whereas alanine, neutrons bound together into a particle identical
leucine, methionine, lysine, and glutamate stabi- to a helium nucleus; it is produced in the radioac-
lize them. Protein alpha-helical regions often fold tive process called alpha decay. Alpha particles,
into coil-coiled configurations that can span like helium nuclei, have a net spin of zero.
membranes, bind DNA, or serve structural roles. The energy of alpha particles varies, depending
upon the specific decay reaction, with higher-
energy alpha particles being emitted from larger
History nuclei, but most alpha particles have energies of
between 3 and 7 MeV, corresponding to
The term alpha helix was coined by William extremely long to extremely short half-lives of
Astbury in the 1930s. Linus Pauling worked out alpha-emitting ▶ nuclides. They are a highly ion-
the structure accurately in 1948. izing form of particle radiation that when
resulting from radioactive alpha decay have low
See Also penetration depth. Helium nuclei, which form
10–12 % of cosmic rays, are usually of much
▶ Amino Acid higher energy than those produced by radioactive
▶ Oligopeptide decay.
▶ Peptide
▶ Polypeptide
▶ Protein
▶ Proteins, Secondary Structure See Also
▶ Beta Rays
▶ Gamma Rays
▶ Radiochemistry
Alpha Particles
▶ Alpha Rays
Alteration
Nicholas Arndt
Alpha Rays ISTerre, Université Grenoble Alpes, France
Jun-Ichi Takahashi
NTT Microsystem Integration Laboratories,
Atsugi, Japan Definition
Alteration in geochemistry refers to processes by

Synonyms which the mineralogy, composition, and texture
of a rock are changed as a result of
Alpha particles; Helium nuclei re-equilibration under conditions of lower tem-
perature and pressure or through interaction with
aqueous or CO2-rich fluids. The minerals of the
Definition original rock, which may be magmatic, sedimen-
tary, or metamorphic, are transformed into an
An alpha ray is a stream of alpha particles. An assemblage of low-temperature, usually finer-
alpha particle consists of two protons and two grained minerals. A typical example is the
al-Tūsı̄, Nasir al-Dı̄n 71
replacement of magmatic minerals such as oliv- Tūsī studied mathematics, medicine, and
ine, pyroxene, and feldspar by chlorite, clay min- Avicennan philosophy at Nishapur, then jurispru-
erals, or carbonates. ▶ Weathering is a type of dence, mathematics, and astronomy in Iraq. In A
alteration that takes place close to the surface 1233, he entered the service of the Shiite Ismaili
through interaction of rock with the atmosphere emir, Ibn Abī Manṣūr, for whom he dedicated his
and with ground- or surface waters. Alteration is work on ethics (Akhlaqi Naṣiri). He spent time in
also used in chemistry and biology (e.g., DNA the Nizari strongholds of Alamut and
alterations). Maymundiz, where he had access to their
libraries and produced a number of his known
works. In 1255, he was sent by the lord of Alamut
to negotiate with Hϋlegϋ Khan (r. 1256–1265),
See Also who led the Mongol westward advances.
He remained in Hϋlegϋ’s service, which may
▶ DNA Damage have been put him in charge of Muslim
▶ Weathering endowments (waqfs) or other important
financial institutions. Hϋlegϋ patronized Tūsī’s
works and allowed him to start the construction
of the Maragha Observatory in 1259. Maragha
became a destination for many scholars who
al-Tūsı̄, Nasir al-Dı̄n
benefitted from Mongol patronage and from the
impressive library that was culled from con-
1 quered libraries in Mesopotamia, Baghdad, and
Harvard Divinity School, Cambridge, MA, USA
2 Syria. Tūsī and his colleagues at Maragha com-
Harvard University, Cambridge, MA, USA
pleted a zij (table of parameters for calculating
the positions of the Sun, Moon, and planets; the
Ilkhanid Zij) around 1270 under Hϋlegϋ’s suc-
Keywords
cessor, Abaqa Khan (r. 1265–1282). The zij
might have been one of the more practical tasks
Islam; Islamic astronomy; Maragha; Medieval;
of the observatory.
Mongol; Zij; Observatory; Tusi couple;
Tūsī is best known for the Tusi couple – a
Avicenna
mathematical device that transforms circular to
linear motion. By rolling a circle of radius r along
the inner edge of a circle of radius 2r, such
Overview that the circles remain tangent and the smaller
circle completes two rotations for every
Nasīr al-Dīn Tūsī (d. 1274) was a Persian scholar rotation of the larger, a given point on the smaller
of mathematics, astronomy, philosophy, and the- circle appears to oscillate along the diameter of
ology; he was considered by many “the Third the larger. The couple was produced as part of
Teacher” after Aristotle and al-Farabi (d. 950). Tūsī’s attempts to address some of the philosoph-
Tūsī’s most renowned activities took place in the ical difficulties of the Ptolemaic model of
Maragha Observatory in what is now Iran, where planetary movements. Applied to the orbs, the
he led a group of scholars including Chinese couple could potentially allow for the observed
astronomers in different investigations and activ- variable length of the orbital radius without
ities. It has been suggested that Tūsī’s criticisms compromising the principle of uniform circular
of Ptolemaic astronomy influenced Copernicus’ movement around the earth. Tūsī preferred this
rejection of equants in his De Revolutionibus solution to Ptolemy’s equant, which generated
(1543). much opposition. Tūsī presented his couple not
72 Aluminilite
as a definitive solution but rather as “an

indication of a solution,” signaling perhaps Aluminilite
that this model was only the beginning of a
longer investigation to be carried out by his ▶ Alunite
students and commentators. While an ostensible
follower of Avicennan philosophy, Tūsī
appeared to have taken a less radical approach
to dealing with the Ptolemaic model than
Avicenna (d. 1037) and his student al-Juzjānī Alunite
(d. 1070), preferring to maintain the model for
its practical values and attempting to Daniele L. Pinti
present important modifications that would GEOTOP Research Center for Geochemistry and
reconcile its contradictions, as opposed to Geodynamics, Université du Québec à Montréal,
rejecting the model for one that would conform Montréal, QC, Canada
better with laws of classical natural
philosophy – a task that was never accomplished
satisfactorily.
Synonyms
In his Ethics, Tusi presented ideas about the
evolution or “perfection” of species, although his
Aluminilite
arguments had important ethical, theological, and
mystical underpinnings.
Definition
Alunite is a secondary mineral of chemical
Dabashi H (1996) The philosopher/vizier: Khwaja Nasir
formula KAl3(SO4)2(OH)6 (trigonal crystal
Al-Din Al-Tusi and the Isma’ilis. Mediaeval Isma’ili
history and thought. Cambridge University Press, system). It forms solid solutions with ▶ jarosite
Cambridge, pp 231–245 KFe3+(SO4)2(OH)6, and it is the product of
Langermann YT (1997) Arabic cosmology. Early Sci Med medium-temperature (80–150 C) hydrothermal
2:185–213
alteration of feldspar-rich volcanic rocks.
Ragep FJ (1987) The two versions of the Tusi couple. In:
King D, Saliba G (eds) From deferent to equant: a vol- Acid fluids formed during the oxidation
ume of studies in the history of science in ancient and and leaching of metal sulfides commonly
medieval near east in honor of E. S. Kennedy, control the alteration. Detection of alunite at
vol 500, Annals of the New York Academy of Sciences.
Terra Sirenum, ▶ Mars, could be an indicator of
New York Academy of Sciences, New York
Ragep FJ (2007) Copernicus and his Islamic predecessors: basalt alteration in contact with H2SO4-rich
some historical remarks. Hist Sci 45:65–81 water. Alunite-▶ kaolinite-layered deposits
Saliba G (1986) The determination of new planetary detected at Columbus Crater are another indica-
parameters at the Maragha observatory. Centaurus
tion of weathering processes at the surface of
29(4):249–271
Saliba G (1995a) A history of Arabic astronomy: plane- ▶ Mars.
tary theories during the golden age of Islam. New York
University Press, New York
Saliba G (1995b) The original source of Qutb Al-Din
Al-Shirazi’s planetary model. In: A history of Arabic
astronomy: planetary theories during the golden age of See Also
Islam, vol 19. New York University Press, New York,
p 119 ▶ Hydrothermal Environments
Tūsi Muḥammad Ibn Muḥammad Naṣīr Al-Dīn, Ragep FJ
˙ (1993) Nasir Al-Din Al-Tusi’s Memoir on Astronomy: ▶ Jarosite
Al-Tadhkira Fīı́lm Al-Hay’a. Springer, New York/ ▶ Mars
Berlin/Paris ▶ Weathering
Amide 73
relatively rare. At this point, the neutral atoms

Amazonian can move relative to the ions, which are effec-
tively tied to the magnetic field. Ambipolar dif- A
Ernst Hauber fusion is thought to occur in ▶ molecular clouds,
Deutsches Zentrum f€ur Luft- und Raumfahrt which are dense enough to shield much of the
(DLR) e.V., Institut f€ur Planetenforschung, external, ionizing radiation. The cloud’s self-
Berlin, Germany gravity can then cause the gas to condense, in
spite of its internal magnetic field. This conden-
sation ultimately leads to star formation.
Definition
See Also
It is the youngest of the three systems (of time-
stratigraphic units) or periods (the chronologic ▶ Fragmentation of Interstellar Clouds
equivalents to systems) in the Martian strati- ▶ Gravitational Collapse, Stellar
graphic scheme, named after the region of ▶ Molecular Cloud
Amazonis Planitia (Amazonis: from the classical ▶ Star Formation, Theory
land of the Amazons on the island Hesperia; see
US Geological Survey Gazetteer of Planetary
Nomenclature). Depending on the different
models to determine absolute ages on planetary Amide
surfaces by crater statistics, the Amazonian
began at some point in time between 3.55 and Henderson James (Jim) Cleaves II
1.8 billion years ago and lasts until the present. Earth–Life Science Institute (ELSI), Tokyo
See Also Japan
▶ Chronostratigraphy USA
▶ Hesperian Blue Marble Space Institute of Science,
▶ Mars Washington, DC, USA
▶ Mars Stratigraphy Center for Chemical Evolution, Georgia Institute
▶ Noachian of Technology, Atlanta, GA, USA
Definition
Ambipolar Diffusion
In chemistry an amide is an organic compound
which contains the functional group or the name
Steven W. Stahler
given to a type of bond formed from the conden-
sation of a carboxylic acid and an ▶ amine.
Monosubstituted amides may exhibits the reso-
nance shown in Fig. 1. Some important amides
Definition
O O−
Ambipolar diffusion is the slippage of neutral R2
matter in a plasma with respect to an internal R2
N N
magnetic field. This slippage occurs when the R1 H R1
H+
ionization fraction is so low that collisions
between neutral species and ions become Amide, Fig. 1 Two states of amide
74 Amidocyanogen
include peptides, urea, and formamide. Amides an amino group has a lone electron pair, amines
are also important intermediates in the Strecker are Lewis bases. Amines are classified as primary
amino acid synthesis. Hydrogen bonding between amines, secondary amines, or tertiary amines
amide functional groups in polypeptides allows depending on the number of alkyl substituents
the formation of secondary structural motifs such (primary amines having a single alkyl substitu-
as a-helices and b-sheets. Amides can be hydro- ent) (Fig. 1). The simplest amine is methylamine
lyzed back to the constituent amine and carboxylic (CH3NH2). Methylamine was found as an inter-
acid. Cyclic amides are known as lactams. stellar molecule in 1974. Amines have also been
detected among organic compounds extracted
from carbonaceous ▶ chondrites.
See Also
See Also
▶ Amine
▶ Carboxylic Acid
▶ Amino Acid
▶ Polypeptide
▶ Chondrite
▶ Strecker Synthesis
▶ Molecular Cloud
Amidocyanogen
Amino Acid
▶ Cyanamide
Jeffrey Bada
CA, USA
Amidogen
▶ Amino Radical Keywords
Amino group; Carboxyl group

Amine
Synonyms
Kensei Kobayashi
Yokohama National University, Tokiwadai,
Amino alkanoic acid
Definition
Definition
Amino acids are organic molecules that contain
An amine is an organic compound containing an
at least one primary amino group (NH2) and one
amino group (-NR3). Since the nitrogen atom in
Amine, Fig. 1 Amine N N N

H H R3
R1 R1 R1
H R2 R2
Primary amine Secondary amine Tertiary amine
Amino Acid 75
Amino Acid, NH2 NH2

Fig. 1 Generalized
structural formulas for a-,
COOH
b-, and g-amino acids
H2N
A
COOH COOH
α-Amino-n-butyric β-Amino-n-butyric γ-Amino-n-butyric

acid acid acid
Amino Acid, Table 1 The numbers of possible structural Amino Acid, Table 2 The names of the structural iso-
isomers for amino alkanoic acids (with the formula mers for 2, 3, 4, and 5 carbon amino alkanoic acids
CnH2nNH2COOH) (Henze and Blair 1934)
Number of
Number of carbon atoms Number of possible isomers carbon atoms Common names
2 1 2 Glycine
3 2 3 Alanine, b-alanine
4 5 4 a-Amino-n-butyric acid, b-amino-
5 12 n-butyric acid, a-aminoisobutyric
6 31 acid, b-aminoisobutyric acid,
g-amino-n-butyric acid
10 1,479
5 Valine, isovaline,
b-aminopentanoic acid,
g-aminopentanoic acid,
carboxyl group (COOH). The general formula for d-aminopentanoic acid,
amino acids with alkyl side chains that have one a-methyl-b-aminobutyric acid,
allo-a-methyl-b-aminobutyric
amino and one carboxyl group, known as amino acid, a-methyl-g-aminobutyric
alkanoic acids, is CnH2nNH2COOH. acid, b-methyl-b-aminobutyric
acid, b-methyl-g-aminobutyric
acid, a-ethyl-b-aminoproponic
acid, a-dimethyl-b-aminoproponic
History acid
Most of the biologically important amino acids

were isolated and characterized in Europe in the Some amino acids have aromatic side chains:
early nineteenth century (Vickery and Schmidt examples are phenylglycine (a-aminophenylacetic
1931). For example, asparagine, the first amino acid), phenylalanine (a-amino-b-phenylpropanoic
acid discovered, was isolated from asparagus by acid), and tyrosine (a-amino-b-(4-hydroxyphenyl)
Vauquelin and Robiquet in 1806. Glycine was propanoic acid). Amino acids can also have side
isolated by Braconnot in 1820. Laboratory syn- chains consisting of an indole (a benzene ring
theses were developed shortly thereafter. linked to a five-membered nitrogen-containing
pyrrole ring) or an imidazole (five-membered
diunsaturated ring composed of three carbon
Overview atoms and two nitrogen atoms at non-adjacent
positions): two examples of this type of amino
The structural isomers with the amino group on acid found in biochemistry are tryptophan and
the sequential carbon atoms adjacent to the car- histidine, respectively.
boxyl group are called a, b, g, etc. -amino acids. There are also amino acids with hydroxyl- and
Thus, a-amino acids are 2-amino alkanoic acids, sulfur-containing side chains. The common
b-amino acids are 3-amino alkanoic acids, names of some examples of these amino acids
etc. General structural formulae for a, b, and g are serine (a-amino-b-hydroxypropanic acid),
amino acids are shown in Fig. 1 and Table 1. threonine (a-amino-b-hydroxybutanoic acid),
The common names of the amino acid isomers cysteine (a-amino-b-mercaptopropionic acid),
with up to five carbon atoms are given in Table 2. and methionine (a-amino-g-(methylthio)butyric
76 Amino Acid
H2N COOH H2N COOH

H
HOOC N
H
N
HOOC Enantiomers
Proline Pipecolic acid
Amino Acid, Fig. 2 Some cyclic amino acids, proline, L-Isoleucine D-Isoleucine
and pipecolic acid
Diastereomers
acid). Selenium can also substitute for sulfur in
the sulfur-containing amino acids in some
organisms.
Some amino acids have more than one amino H2 N COOH H2N COOH
group (e.g., lysine or a, e-diaminohexanoic acid)
and/or more than one carboxyl group: for exam-
ple, a-aminomalonic acid, aspartic acid Enantiomers
(a-aminobutanedioic acid), and glutamic acid
(a-aminopentanedioic acid). Asparagine
(a-amino-b-carbamoylpropanoic acid) and gluta-
mine (a-amino-d-carbamoylbutyric acid) are the L-Alloisoleucine D-Alloisoleucine
side group carboxamides of aspartic and glutamic Amino Acid, Fig. 3 Enantiomers and diastereomers of
acids, respectively. Interestingly, asparagine was iso- and alloisoleucine
the first amino acid discovered in 1806 when it
was crystallized from the “juice” squeezed from 3.9 at 25 C; thus at neutral pH aspartic acid has a
asparagus shoots. The amino acid arginine net negative charge. The pKa of the guanidinium
(a-amino-e-guanidinopentanoic acid) has a group of arginine is 12.5, and arginine is thus
guanidinium group attached to the end of its positively charged at neutral pH.
alkyl side chain. When a carbon atom in an amino acid has four
Some amino acids have a cyclic secondary different groups attached to it, referred to as an
amine rather than a primary amino group. Exam- asymmetric or ▶ chiral carbon, it is optically
ples include proline (pyrrolidine-2-carboxylic active. For those amino acids with one chiral
acid, C5H9NO2) where the primary amino group carbon, there are two possible optically active
is replaced with a five-membered pyrrolidine ring isomers designated, the L- and D-enantiomers.
or tetrahydropyrrole and pipecolic acid Some amino acids have more than one chiral
(piperidine-2-carboxylic acid, C6H11NO2) carbon so several stereoisomers are possible. An
where the amino group is replaced by amino acid with two chiral carbons is said to be
six-membered piperidine ring (Fig. 2). diastereomeric, and there are thus two diastereo-
The ionization constants (pKa) at 25 C of the mers, each of which has two enantiomers, for a
amino and carboxyl groups of amino alkanoic total of four possible optical isomers. For the
acids are in the range 8–10 and 2–4, respectively. diastereomeric pair L-isoleucine/D-alloisoleucine
Thus, at neutral pH, the amino group is proton- (a-amino-b-methylpentanoic acid), the two sets
ated, while the carboxyl group is deprotonated, of enantiomers are L- and D-isoleucine and L- and
producing a doubly charged ▶ zwitterion with no D-alloisoleucine, respectively (Fig. 3).
net charge. Amino acids with other amino or Amino acids can be linked together by the
carboxyl groups have additional ionization conformation of a ▶ peptide bond that involves the
stants characteristic of the particular group. The amino group of one amino acid and the carboxyl
pKa of the b-carboxyl group of aspartic acid is group of another. Two amino acids connected in
Amino Acid 77
O R Amino Acid, Table 3 The 20 amino acids commonly

found in proteins and their commonly used abbreviations
+ Amino acid Three-letter One-letter
H3N
common name abbreviation abbreviation
A
N −
COO
H Aspartic acid Asp D
Glutamic acid Glu E
Amino Acid, Fig. 4 A generic dipeptide
Asparagine Asn N
Glutamine Gln Q
Glycine Gly G
this fashion are called a dipeptide which has the
Alanine Ala A
structure given below (in this case, one amino
Valine Val V
acid is glycine and the other has a generic Isoleucine Iso I
R-group side chain) (Fig. 4). Leucine Leu L
Polypeptides and proteins consist of a large Phenylalanine Phe F
number of amino acids connected in peptide link- Tyrosine Tyr Y
ages. A total of 20 different amino acids (for this Serine Ser S
discussion, the amino acids selenocysteine and Threonine Thr T
pyrrolysine are not included because they are rel- Cysteine Cys C
atively rare coded amino acids) are decoded from Methionine Met M
DNA sequences and encoded into RNA for incor- Lysine Lys K
poration into proteins. A list of the “canonical” Histidine His H
20 protein amino acids and their abbreviations are Arginine Arg R
given in Table 3. There are peptides that contain Tryptophan Trp W
additional amino acids other than the standard Proline Pro P
protein amino acids, but these are incorporated
by posttranslational modifications or the peptides
themselves are synthesized by nonribosomal pep- meteorites, and there are hints that at least the
tide synthetases (NRPSs). For example, simplest amino acid glycine is present in interstel-
a-aminoisobutyric acid and isovaline are found lar clouds and comets (see ▶ Molecules in Space).
in some fungal peptides synthesized by NRPSs. One of the meteorites most extensively studied is
With the exception of achiral glycine, only the the ▶ Murchison carbonaceous chondrite that fell
L-enantiomers of the proteinogenic amino acids are in southeastern Australia in 1969. Over 75 differ-
incorporated into proteins. The discrimination ent amino acids have been detected in Murchison
against the incorporation of D-amino acids during (Sephton 2002), with only 8 of these also being
the protein synthesis process is estimated to be found in biological proteins. These amino acids
greater than 104. However, there are D-amino acids are clearly of extraterrestrial origin: many are
present in some peptides, but these are introduced unique to the meteorite and do not occur naturally
either by the conversion of L-amino acids by post- on Earth and those with a chiral carbon are race-
translational isomerization enzymes or are intro- mic (or close to racemic). The Murchison amino
duced by NRPSs. Some D-amino acid-containing acids are thought to have been synthesized by
peptides have potent antimicrobial activity. natural reactions, such as the ▶ Strecker synthesis,
The total number of amino acids theoretically directly on the juvenile meteorite parent body or in
possible is huge, and several hundred different the early solar nebula before incorporation into
amino acids have been isolated from organisms, planetesimals. Amino acids have been detected
and an even larger number have been made in the in even larger quantities in other carbonaceous
laboratory by a variety of synthetic methods. chondrites (Pizzarello and Shock 2010).
Moreover, the synthesis of amino acids is not Amino acids may also have been synthesized by
confined to terrestrial biology or laboratory syn- natural processes on the early Earth as demon-
thesis: amino acids have been detected in strated by the classic Miller spark discharge
78 Amino Acid N-Carboxy Anhydride
experiment carried out in 1953 (Miller 1953; John- ▶ L-Amino Acids

son et al. 2008). These amino acids could have ▶ Molecules in Space
accumulated on the Earth and been available for ▶ Peptide
incorporation into the first living entities. To date, ▶ Protein
12 of the amino acids found in the proteins of ▶ Strecker Synthesis
terrestrial organisms have been synthesized in
spark discharge experiments with various reduced
gas mixtures.
The striking overlap between amino acids Cleaves HJ (2010) The origin of the biologically coded
generated in experiments simulating prebiotic amino acids. J Theor Biol 263:490–498
chemistry and found in meteorites represents evi- Henze HR, Blair CM (1934) The number of structural
dence for the abiotic plausibility of approxi- isomers of the more important types of aliphatic com-
pounds. J Am Chem Soc 56:157
mately half of the canonical amino acid set. Higgs PG, Pudritz RE (2009) A thermodynamic basis for
This is also supported by the thermodynamics of prebiotic amino acid synthesis and the nature of the
their formation (Higgs and Pudritz 2009). Debate first genetic code. Astrobiology 9:483–490
continues, however, as to whether these amino Johnson AP, Cleaves HJ, Dworkin JP, Glavin DP,
Lazcano A, Bada JL (2008) The Miller volcanic
acids would have been sufficient to comprise the spark discharge experiment. Science 322:404
first functional proteins. It is also unclear how Miller SL (1953) Production of amino acids
these amino acids came to be used by life when under possible primitive Earth conditions. Science
many others might have been also available 117:528
Philip GK, Freeland SJ (2011) Did evolution select a
(Weber and Miller 1981; Cleaves 2010). nonrandom “alphabet” of amino acids? Astrobiology
The amino acids absent from abiotic simula- 11(3):235–240
tions and meteorites are not only difficult to make Pizzarello S, Shock E (2010) The organic composition of
but are also either thermally unstable (Gln, Asn) or carbonaceous meteorites: the evolutionary story ahead
of biochemistry. In: Deamer D, Szostak J (eds) The
are unstable under UV conditions (Cys, Met, Trp, origins of life. Cold Spring Harbor Press, Washington,
His, Tyr, Phe). This strongly suggests that these DC, pp 89–107
amino acids are biological “inventions” and their Sephton MA (2002) Organic compounds in carbonaceous
multi enzymatic synthesis pathways appeared only meteorites. Nat Prod Rep 19:292–311
Vickery HB, Schmidt CLA (1931) The history of the
after the onset of Darwinian evolution. The order discovery of the amino acids. Chem Rev 9(2):169–318
of biologically invented amino acid entering the Weber AL, Miller SL (1981) Reasons for the occurrence
code has been hypothesized based on thermody- of the twenty coded protein amino acids. J Mol
namics (Higgs and Pudritz 2009); however, the Evol 17:273–284
entry and retention of amino acids is likely to be
complex, with such factors as accessibility, com-
patibility, complementarity with existing code
members, and stability on transfer RNA all
playing a role (Weber and Miller 1981; Cleaves Amino Acid N-Carboxy Anhydride
2010). The culmination of this selection is a set of
amino acids unchanged since the last universal Laurent Boiteau
common ancestor (LUCA), which is exceptional Institut des Biomolécules Max Mousseron,
in breadth and evenness in terms of size, hydro- UMR5247 CNRS, University Montpellier-2,
phobicity, and charge (Philip and Freeland 2011). Montepellier, Cedex, France
See Also Synonyms
▶ Diastereomers 1,3-Oxazolidine-2,5-dione; Leuchs’ Anhydride;

▶ Enantiomers NCA
Amino Acid N-Carboxy Anhydride 79
Definition emergence of homochirality of the natural amino

acid pool, e.g., enantiomeric excess amplification
An amino acid N-carboxy anhydride (or NCA) is a processes (Kricheldorf 2006; Pascal et al. 2005; A
cyclic organic compound structurally related to an Illos et al. 2008). NCAs have long been postulated
▶ amino acid, which is an intramolecular mixed as likely intermediates in the reaction of activated
anhydride of a carboxylic and carbamic acid amino acid esters (e.g., adenylates, thioesters)
(structure -CO-O-CO-NH-), making it both an based on the observation that the formation of
N-protected and a CO-activated amino acid. NCAs peptides is accelerated by the presence of CO2 or
are structurally related to hydantoins but have very bicarbonate. Since the late 1990s, several
different chemical reactivity. They are rather unsta- prebiotically relevant pathways for NCA forma-
ble in water and physiological media. The term tion have been identified, thus confirming the pre-
NCA is usually used to refer to the NCAs of biotic status of NCAs (Kricheldorf 2006; Pascal
a-amino acids (although NCAs of b-amino acids et al. 2005):
etc. are also possible). NCAs can condense to give
oligo- or polypeptides, with the release of CO2. • The nitrosation of N-carbamoyl amino acids
NCAs are postulated or observed intermediates in (CAA) promoted by nitrogen oxides
many prebiotically relevant reactions leading to (Kricheldorf 2006; Pascal et al. 2005)
▶ peptides from amino acid derivatives, especially • The decomposition of diacyldisulfides
in aqueous media in the presence of carbonate. • The reaction of amino acids with carbonyl
NCAs are also considered to be potentially prebiotic sulfide in the presence of oxidizing or
reagents, as they are versatile free energy carriers alkylating agents (Leman et al. 2004)
which can potentially activate other biologically • The spontaneous decomposition of CAA in
relevant chemical species, such as nucleotides. water (Danger et al. 2006)
NCAs represent both (1) the structurally sim-

History plest activated amino acids (formally resulting
from condensation with CO2), (2) an unavoidable
Although speculations that NCAs might have intermediate from any form of CO-activated amino
played a role in prebiotic chemical evolution acid in a bicarbonate/CO2-rich environment, and
arose in the mid-1970s, notwithstanding their use (3) the most activated amino acid species achiev-
since the late 1970s in “model” prebiotic reactions, able in water in a prebiotic environment. Thermo-
studies in the early 2000s improved the status of dynamic calculations show NCAs to be quite stable
NCAs as prebiotically relevant compounds. (because of the cyclic structure) compared to other
anhydrides, although kinetically they are as reac-
tive as the latter. Furthermore, NCAs may be kinet-
Overview ically competent intermediates from almost any
inactivated amino acid derivatives, provided their
Discovered by Hermann Leuchs in 1906, NCAs spontaneous hydrolysis is slower than NCA forma-
are well-known reactants in both organic and tion (Pascal et al. 2005).
polymer synthesis (Kricheldorf 2006). Since Such thermodynamic and kinetic features make
their most popular preparative method involving NCAs potential energy carriers in an amino acid-
the reaction of free amino acids with phosgene is based protometabolism, as exemplified by their
not prebiotically relevant, NCAs themselves were ability to activate inorganic phosphate (Pascal
long considered as prebiotically irrelevant (Pascal et al. 2005) or nucleotides (Biron et al. 2005;
et al. 2005). Nevertheless, NCAs have been con- Leman et al. 2006), which could be coupled to a
tinuously used from the 1970s in model reactions peptide/nucleic acid coevolution scenario
of prebiotic peptide formation, especially to assess supporting speculations on the emergence of the
stereoselection hypotheses in relation with the translation apparatus (Pascal et al. 2005).
80 Amino Acid Precursors
See Also hydrolysis). One of the typical amino acid precur-

sors is ▶ aminoacetonitrile, which is converted to
▶ Amino Acid glycine by hydrolysis via glycine ▶ amide:
▶ Chirality NH2CH2CN + 2H2O ! NH2CH2CONH2 + H2O
▶ Metabolism, Prebiotic ! NH2CH2COOH + NH3. Hydantoins
▶ N-Carbamoyl Amino Acid (substituted glycolylurea) are also typical amino
▶ Peptide acid precursors and have been found in carbona-
▶ Prebiotic Chemistry ceous chondrites. Complex organic polymers with
large molecular weights are also possible precur-
sors. ▶ Tholins, which are formed by reactions of
mixtures of nitrogen and methane, are large com-
Biron JP, Parkes AL, Pascal R, Sutherland JD plex molecules which give amino acids after
(2005) Expeditious prebiotic aminoacylation of nucle- hydrolysis. Amino acids are frequently detected
otides. Angew Chem Int Ed 44:6731–6734 in carbonaceous ▶ chondrites (meteorites), but the
Danger G, Cottet H, Boiteau L, Pascal R (2006) The amount of amino acids recovered usually
peptide formation mediated by cyanate revisited.
N-Carboxyanhydrides as accessible intermediates in increases after hydrolysis, suggesting that some
the decomposition of N-carbamoylamino acids. J Am amino acids are present in the form of amino
Chem Soc 128:7412–7413 acid precursors.
Illos RA, Bisogno FR, Clodic G, Bolbach G, Weissbuch I,
Lahav M (2008) Oligopeptides and copeptides of
homochiral sequence, via b-sheets, from mixtures of
racemic a-amino acids, in a one-pot reaction in water; See Also
relevance to biochirogenesis. J Am Chem Soc
130(27):8651–8659 ▶ Amide
Kricheldorf HR (2006) Polypeptides and 100 years of
chemistry of a-amino acid N-carboxyanhydrides. ▶ Amino Acid
Angew Chem Int Ed 45:5752–5784 (and references ▶ Aminoacetonitrile
cited therein) ▶ Chondrite
Leman L, Orgel LE, Ghadiri MR (2004) Carbonyl sulfide- ▶ Complex Organic Molecules
mediated prebiotic formation of peptides. Science
306:283–286 ▶ Hydantoin
Leman LJ, Orgel LE, Ghadiri MR (2006) Amino acid ▶ Hydrolysis
dependent formation of phosphate anhydrides in ▶ Tholins
water mediated by carbonyl sulfide. J Am Chem Soc
128(1):20–21
Pascal R, Boiteau L, Commeyras A (2005) From the prebi-
otic synthesis of a-amino acids towards a primitive
translation apparatus for the synthesis of peptides. Top
Curr Chem 259:69–122 (and references cited therein) Amino Alkanoic Acid
▶ Amino Acid
Amino Acid Precursors
Kensei Kobayashi Amino Radical

Hodogayaku, Yokohama, Japan William M. Irvine
Definition
Synonyms
▶ Amino acid precursors are compounds that
give amino acids after some reactions (usually Amidogen; Aminyl radical; NH2
Aminoacetonitrile 81
Definition
Aminoacetonitrile
This triatomic radical is an important A
intermediary in the interstellar chemistry of Didier Despois
▶ ammonia, NH3. Like many light hydrides, its Laboratoire d’Astrophysique de Bordeaux,
pure rotational transitions occur at far infrared/ CNRS-Universite de Bordeaux, France
submillimeter wavelengths, making its observa-
tion from ground-based observatories difficult
because of the opacity of the terrestrial Synonyms
atmosphere.
AAN; Cyanomethylamine; Glycinonitrile;
NH2CH2CN
History
The NH2 radical was first detected in the ▶ inter- Definition

stellar medium in 1993 (van Dishoeck et al.), at
submillimeter wavelengths, and has been Aminoacetonitrile (IUPAC name 2-
extensively observed toward different Aminoacetonitrile) is a (toxic) liquid at room tem-
molecular clouds using the HIFI instrument on perature and standard pressure. It is a precursor of
board the Herschel satellite (Persson et al. 2010, the simplest amino acid, ▶ glycine, which it forms
2012). by reaction with liquid water. It is also an interme-
diary in the ▶ Strecker synthesis of glycine. It was
identified in the interstellar medium in 2008.
See Also
▶ Ammonia History
▶ Interstellar Medium
▶ Molecules in Space Although its rotational spectrum has been studied
since the 1970s, and modeled explicitly for a
search in the interstellar medium in 1990,
aminoacetonitrile has only been detected recently
References and Further Reading in space, in a large molecular cloud Sagittarius
B2 (Sgr B2) at the center of the Galaxy (Belloche
Persson CM, Black JH, Cernicharo J et al (2010) Nitrogen et al. 2008).
hydrides in interstellar gas. Herschel/HIFI observa-
tions towards G10.6-0.4 (W31C). Astron Astrophys
521:L45 See Also
Persson CM, De Luca M, Mookerjea B et al (2012) Nitro-
gen hydrides in interstellar gas. II. Analysis of Her-
schel/HIFI observations towards W49N and G10.6-0.4 ▶ Glycine
(W31C). Astron Astrophys 543:145 ▶ Molecular Cloud
van Dishoeck EF, Jansen DJ, Schilke P, Phillips TG ▶ Molecules in Space
(1993) Detection of the Interstellar NH2 Radical. ▶ Strecker Synthesis
Astrophys J Lett 416:L83–L86
Belloche A, Menten KM, Comito C, M€ uller HSP,

Aminoacetic Acid Schilke P, Ott J, Thorwirth S, Hieret C (2008) Detec-
tion of amino acetonitrile in Sgr B2(N). Astron
▶ Glycine Astrophys 482:179–196
82 Aminobutyric Acid
See Also
Aminobutyric Acid
▶ Amino Acid
Mark Dörr
University of Southern Denmark, Odense M,
Denmark
Aminoethanoic Acid
Synonyms
▶ Glycine
Butyrine; Ethyl-glycine
Definition Aminoisobutyric Acid
Aminobutyric acid is the term for a variety of Henderson James (Jim) Cleaves II
structural isomers of amino acids derived from n- Earth–Life Science Institute (ELSI), Tokyo
or isobutyric acid with the chemical formula Institute of Technology, Meguro–ku,
C4H9NO2. They belong to the substance class of Tokyo, Japan
amino acids, since they contain an amino func- Institute for Advanced Study, Princeton, NJ,
tional group and a carboxylic acid functional USA
group. In nature, several different isomers of Blue Marble Space Institute of Science,
aminobutyric acid are found: (1) a-aminobutyric Washington, DC, USA
acid (aABA), a key intermediate in the biosynthe- Center for Chemical Evolution, Georgia Institute
sis of ophthalmic acid, (2) b-aminobutyric acid of Technology, Atlanta, GA, USA
(bABA), (3) g-aminobutyric acid (GABA),
which modulates the excitability of neurons of Synonyms
vertebrates and muscle tone, and
(4) a-aminoisobutyric acid (aAIB), which is AIB
found in some fungal membrane peptides. Several
aminobutyric acid isomers have been found in
carbonaceous chondrite meteorites. Definition
a, b, and g denote the position of the amino
group relative to the carboxyl group in the Amino isobutyric acid (AIB) is an amino acid
▶ amino acid molecule: a refers to the first, b derived from isobutyric acid. There are two struc-
the second, and g the third position. tural isomers of amino isobutyric acid (Fig. 1),
a-aminoisobutyric acid (aAIB), which is achiral,
O
and b-aminoisobutyric acid (bAIB), which has
NH2 O
two ▶ stereoisomers, a D and L form. Both
α
OH
β OH H2N COOH COOH
NH2
α-amino butyric acid β-amino butyric acid
O
H2N
α AIB β AIB
H2N γ
OH
Aminoisobutyric Acid, Fig. 1 The two structural iso-
γ-amino butyric acid mers of aminoisobutyric acid
Amitsoq Gneisses 83
isomers have been found in carbonaceous chon- See Also

drites, with aAIB often being one of the most
abundant amino acids. This is thought to be sig- ▶ Amino Acid A
nificant as aAIB is not found in proteins, ▶ Amino Acid Precursors
suggesting an extraterrestrial origin of this com- ▶ Cyanamide
pound. However, several fungi are now known to ▶ Strecker Synthesis
synthesize this compound for incorporation in
non-ribosomally encoded peptide antibiotics.
See Also
Aminyl Radical
▶ Carbonaceous Chondrites, Organic
▶ Amino Radical
Chemistry of
▶ Stereoisomers
Amitsoq Gneisses
Aminonitrile Hervé Martin
Laboratoire Magmas et Volcans, Université
Blaise Pascal, OPGC, CNRS, IRD, Clermont-
Ferrand, France
Institute of Technology, Meguro–ku,
Tokyo, Japan
Keywords
USA
Greenland; Isua Supracrustal Belt; Archean;
Washington, DC, USA
TTG; Gneiss; Metamorphic rocks
Synonyms
Definition
Itsaq Gneiss Complex
An aminonitrile is a compound containing both
an amino and a nitrile functional group. The
simplest aminonitrile is ▶ cyanamide. Definition
a-Aminonitriles, such as a-aminoacetonitrile,
are important intermediates in the ▶ Strecker The Amı̂tsoq gneisses are among the older meta-
synthesis of amino acids, as they are hydrolyzed morphic rock complexes yet discovered on Earth.
consecutively to a-amino amides and finally to These rocks outcrop on the southwestern coast of
a-amino acids (Fig. 1). a-Aminoacetonitrile was Greenland, where they extend over more than
detected in interstellar space in 2008. 50 km northeast of Nûk (Godthåb). The oldest
O NH4CN
Aminonitrile, H2N CN H2O H2N CONH2 H2O H2N COOH
Fig. 1 Strecker amino acid
synthesis via aminonitrile R1 R2 R2 R1 R2 R1 R2 R1
α-amino nitrile α-amino amide α-amino acid
84 Amitsoq Gneisses
age obtained on a zircon crystal extracted from a This view has been subsequently corroborated
tonalitic gneiss sample is of 3.872 0.010 Ga. by intensive zircon dating (Nutman et al. 1996;
Nutman and Hiess 2009) that determined that the
emplacement of Amı̂tsoq gneiss protolith
Overview (protolith is the original, unmetamorphosed rock
from which a metamorphic rock is formed)
The Amı̂tsoq gneisses outcrop on the southwest- occurred between 3.88 and 3.60 Ga, during at
ern coast of Greenland, where they extend over least three main petrogenetic episodes at
more than 50 km northeast of Nûk (Godthåb), ~3.80 Ga; ~3.7 Ga, and ~3.65 Ga. It must be
along the southern coast of the Godthåbsfjord. noted that some ages older than 3.85 Ga were
After long and detailed field work, Mc Gregor also measured (Horie et al. 2010; Nutman
(1968, 1973) was the first who recognized these et al. 2013). Indeed, a zircon crystal extracted
Archaean terrains as among the oldest in the from a tonalitic gneiss sample gave an age of
world. He distinguished two groups of gneisses: 3.872 0.010 Ga, which is the oldest reliable
(1) very old ones cut by mafic dykes (Ameralik age so far measured in Amı̂tsoq gneisses. The
dykes) that he called the Amı̂tsoq gneisses, same authors reported an age of
(2) younger ones emplaced after the Ameralik 3.883 0.009 Ga measured in a zircon core,
dykes and that are known as the Nûk gneisses. while the rim provided a slightly younger age of
Both groups are crosscut by the late Qôrqut gran- 3.861 0.022 Ga; these dates are assumed to be
ite. The first dating of the Amı̂tsoq gneisses was those of parental magma crystallization.
conducted by Black et al. (1971), who obtained a The Amı̂tsoq gneisses outcrop over vast areas,
Rb-Sr isochron age of 3.98 0.17 Ga, and over about 3,000 km2. Their protolith was a felsic
Moorbath et al. (1972), who measured a slightly plutonic rock, having a tonalitic, trondhjemitic,
younger age (3.74 0.1 Ga) using the same and granodioritic (TTG) composition (O’Nions
method. More recent researches showed that the and Pankhurst 1978; Nutman and Bridgwater
so-called Amı̂tsoq gneisses were heterogeneous 1986; Nutman et al. 2000, 2007, 2013; Steenfelt
and made up of several intrusive bodies. In order et al. 2005; Hiess et al. 2009). In Archean ter-
to account for this diversity, Nutman et al. (1996) rains, TTGs are very abundant; these are by far
proposed to refer to these formations as the Itsaq the most abundant rocks of the Archean continen-
Gneiss Complex. In fact, both terms are used in tal crust (Moyen and Martin 2012). They are
geological literature. generated by partial melting of a hydrous basalt,
Amitsoq Gneisses,
Fig. 1 General view of the
~3.8 Ga Amı̂tsoq gneisses.
They consist in grey gneiss,
TTG in composition. On
this photo, they are crosscut
by a black Ameralik dyke
(Photo G. Gruau)
Amitsoq Gneisses 85
possibly in a subduction-like environment Hiess J, Bennett VC, Nutman AP, Williams IS (2009) In
(Martin 1986; Martin et al. 2005). Subsequently, situ U-Pb, O and Hf isotopic compositions of zircon
and olivine from Eoarchaean rocks, West Greenland:
they underwent granulite facies metamorphism at new insights to making old crust. Geochim A
about 3.6 Ga (Friend and Nutman 2005). Cosmochim Acta 73:4489–4516
Before the Cretaceous, and the development Horie K, Nutman AP, Friende CRL, Hidaka H (2010)
of the Labrador ridge and the Baffin Bay basin, The complex age of orthogneiss protoliths
exemplified by the Eoarchaean Itsaq Gneiss Complex
west Greenland and Labrador were closer to each (Greenland): SHRIMP and old rocks. Precambrian Res
other. At that time, Archean terrains on both sides 183:25–43
should have been connected. Indeed, along the Martin H (1986) Effect of steeper Archean geothermal
northern coast of Labrador outcrop the Uivak gradient on geochemistry of subduction-zone magmas.
Geology 14:753–756
gneisses, which are metamorphic rocks, mostly Martin H, Smithies RH, Rapp R, Moyen J-F, Champion
TTG in composition and very similar to the D (2005) An overview of adakite, tonalite-
Amı̂tsoq gneisses. The Uivak gneisses contain trondhjemite-granodiorite (TTG), and sanukitoid:
zircon crystals that have been dated at relationships and some implications for crustal evolu-
tion. Lithos 79:1–24
3.733 0.009 Ga; however, they also contain McGregor VR (1968) Field evidence of very old Precam-
rounded cores dated at 3.863 0.012 Ga brian rocks in Godthaab area, West Greenland.
(Bridgwater and Collerson 1976; Bridgwater Rapp Gronlands Geol Unders 19:31
and Schiøtte 1991). Contrarily to Greenland, in McGregor VR (1973) The early Precambrian geology of
the Godthåb district, West Greenland. Phil Trans
Labrador these rocks suffered a Neoarchean R Soc Lond A 273:243–258
granulite facies metamorphism (Collerson and Moorbath S, O’Nions RK, Pankhurst RJ, Gale NH,
Bridgwater 1979) (Fig. 1). McGregor VR (1972) Further rubidium-strontium
age determinations on the very eary Precambrian
rocks of Godthaab region, West Greenland. Nature
See Also 240:78–82
Moyen J-F, Martin H (2012) Forty years of TTG research.
Lithos 148:312–336
▶ Archean Eon Nutman AP, Bennett VC, Friend CLR, Horie K, Hidaka
▶ Earth’s Atmosphere, History of the Origins H (2007) ~3,850 Ma tonalites in the Nuuk region,
▶ Greenland Greenland: geochemistry and their reworking within
an Eoarchaean gneiss complex. Contrib Mineral Petrol
154:385–408
▶ Metamorphic Rock Nutman AP, Bennet VC, Friend CLR, McGregor VR
▶ Tonalite-Trondhjemite-Granodiorite (2000) The early Archaean Itsaq Gneiss
Complex of southern Greenland: the importance
of field observations in interpreting age
and isotopic constrains for early
References and Further Reading terrestrial evolution. Geochim Cosmochim Acta
64:3035–3060
Black LP, Gale NH, Moorbath S, Pankhurst RJ, McGregor Nutman AP, Bridgwater D (1986) Early
VR (1971) Isotopic dating of very early Precambrian Archaean Amitsôq tonalites and granites of the
amphibolite facies gneisses from the Godthaab district, Isukasia area, southern West Greenland: development
West Greenland. Earth Planet Sci Lett 12:245–259 of the oldest known sial. Contrib Mineral Petrol
Bridgwater D, Collerson KD (1976) The major petrolog- 94:137–148
ical and geochemical characters of the 3600 Nutman AP, McGregor VR, Friend CLR, Bennet VC,
m.y. Uivak gneisses from Labrador. Contrib Mineral Kinny PD (1996) The Itsaq gneiss complex of southern
Petrol 54:43–60 Greenland; the world’s most extensive record of early
Bridgwater D, Schiøtte L (1991) The Archaean gneiss crustal evolution (3900–3600 Ma). Precambrian Res
complex of northern Labrador. A review of current 78:1–39
results, ideas and problems. Bull Geol Soc Den Nutman AP, Bennett VC, Friend CLR, Hidaka H, Yi K,
39:153–166 Ryeol Lee S, Kamiichi T (2013) The Itsaq
Friend CRL, Nutman AP (2005) Complex 3670–3500 Ma Gneiss Complex of Greenland: episodic 3900 to
orogenic episodes superimposed on juvenile crust 3660 Ma juvenile crust formation and recycling in
accreted between 3850–3690 Ma, Itsaq Gneiss Com- the 3660 to 3600 Ma Isukasian orogeny. Am J Sci
plex, southern West Greenland. J Geol 113:375–398 313:877–911
86 Ammonia
Nutman AP, Hiess J (2009) A granitic inclusion suite History

within igneous zircons from a 3.81 Ga tonalite
(W. Greenland): restrictions for Hadean crustal evolu-
tion studies using detrital zircons. Chem Geol Ammonia has been known since ancient times,
261:76–81 although it was first isolated by Priestly in 1774.
O’Nions RK, Pankhurst RJ (1978) Early Archaean rocks In 1785, Berthollet determined its composition.
and geochemical evolution of the Earth’s crust. Earth The Haber-Bosch process to synthesize ammonia
Planet Sci Lett 38:211–236
Steenfelt A, Garde AA, Moyen J-F (2005) Mantle wedge from nitrogen and hydrogen was developed by
involvement in the petrogenesis of Archaean grey Fritz Haber and Carl Bosch in 1909. It was found
gneisses in West Greenland. Lithos 79:207–228 in space by Cheung et al. (1968) and in comets by
Altenhoff et al. (1983) using radioastronomical
techniques.
Ammonia Overview
Alexander Smirnov Ammonia has been detected throughout our solar

Department of Earth and Marine Science, system as well as in interstellar space (see ▶ Mol-
Dowling College, Oakdale, NY, USA ecules in Space). The deuterated ammonium ion
NH3D+ has recently been detected in the inter-
stellar medium (Cernicharo et al. 2013; note that
Keywords the symmetry of the principle isotopic form,
NH4+, leads to a zero electric dipole moment
Prebiotic synthesis; Nitrogen; Abiotic reduction and hence no pure rotational transitions that
might be observed astronomically). Ammonia is
found as a gas in planetary atmospheres and in the
Synonyms solid form (ice) in cometary nuclei and planetary
surfaces. Ammonia is hypothesized to be present
Azane; Nitro-sil; Trihydrogen nitride in liquid form in a subsurface ocean on some
outer planet satellites (e.g., ▶ Titan) where it
would effectively lower the freezing point of
Definition water (Raulin 2008).
On the early Earth, ammonia was likely a
Ammonia (NH3) is a chemical compound com- necessary precursor for prebiotic organic synthe-
posed of ▶ nitrogen and hydrogen which exists as sis, such as the ▶ Strecker synthesis of amino
a gas at standard conditions of temperature acids. It was used as the nitrogen source in the
and pressure. In the trigonal pyramidal ammonia Miller-Urey experiment, which produced a suite
molecule, the lone electron pair of the nitrogen of organic compounds such as amino acids from a
atom is responsible for its dipole moment mixture of reduced gases simulating the primor-
(polarity) and its behavior as a base (proton dial atmosphere (Miller 1953). However, most
acceptor). It dissolves readily in water and its current models suggest the early atmosphere
protonation results in the formation of the conju- was only mildly reducing, with the redox state
gate acid ammonium ion (NH4+) with both spe- linked to the evolution and oxidation state of the
cies coexisting in a pH-dependent equilibrium Hadean and early Archaean mantle, with
(pKa NH4+ = 9.25 at 25 C). Liquid ammonia ▶ dinitrogen (N2) as the dominant nitrogen spe-
(boiling point 33.35 C at atmospheric cies (Kasting and Catling 2003).
pressure) is an ionizing solvent with physical It has been experimentally shown that
properties and behavior similar to water ammonia-containing environments are more effi-
(Lagowski 2007). cient in organic synthesis than those dominated
Ammonium, Deuterated 87
by dinitrogen in both aqueous and gaseous envi- Cernicharo J, Tercero B, Fuente A, Domenech JL,
ronments. This notion is not unexpected, consid- Cueto M, Carrasco E, Herrero VJ, Tanarro I,
Marcelino N, Roueff E (2013) Detection of the ammo-
ering that the strong triple bond (948 kJ.mol1) of nium ion in space. Astrophys J 771:L10–L13 A
the N2 molecule results in large reaction activa- Cheung AC, Rank DM, Townes CH, Thornton DD, Welch
tion energy barriers even if the overall reaction is WJ (1968) Detection of NH3 molecules in the inter-
thermodynamically favored. The process of con- stellar medium by their microwave emission. Phys Rev
Lett 21:1701
version (e.g., reduction) of unreactive dinitrogen Kasting JF, Catling D (2003) Evolution of a habitable
to reactive and prebiologically useful ammonia is planet. Annu Rev Astron Astrophys 41:429–463
referred to as ▶ nitrogen fixation. Lagowski JJ (2007) Liquid ammonia. Synth React Inorg
Mechanisms suggested for abiotic ammonia Met 37:115–153
Miller SL (1953) A production of amino acids under
production on the early Earth include reduction possible primitive Earth conditions. Science
of atmospherically derived nitrite (NO2) by fer- 117:528–529
rous iron or iron-bearing minerals (Summers and Raulin F (2008) Astrobiology and habitability of Titan.
Chang 1993); hydrolysis of atmospherically pro- Space Sci Rev 135:37–48
Singireddy S, Gordon AD, Smirnov A, Vance MA,
duced HCN (Zahnle 1986); reduction of Schoonen MA, Szilagyi RK, Strongin DR
dinitrogen on mineral surfaces (sulfides, metals, (2012) Reduction of nitrite and nitrate to ammonium
alloys) in hydrothermal systems (Brandes on pyrite. Orig Life Evol Biosph 42:275–294
et al. 2008; Smirnov et al. 2008; Singireddy Smirnov A, Hausner D, Laffers R, Strongin D, Schoonen
MA (2008) Abiotic ammonium formation in the pres-
et al. 2012); or delivery of reduced nitrogen ence of Ni-Fe metals and alloys and its implications for
(nitride, N3) in iron meteorites followed by dis- the Hadean nitrogen cycle. Geochem Trans 9:5
solution and reaction with H+ (Smirnov Summers DP, Chang S (1993) Prebiotic ammonia from
et al. 2008). The concentrations of ammonia reduction of nitrite by iron(II) on the early Earth.
Nature 365:630–632
and/or ammonium ion in the prebiotic atmosphere Zahnle K (1986) Photochemistry of methane and the for-
and hydrosphere were likely controlled by mech- mation of Hydrocyanic acid (HCN) in Earth’s early
anisms such as photolytic destruction, sequestra- atmosphere. J Geophys Res 91:2819–2834
tion in clay minerals by substitution for K+ and
formation of N-bearing organic molecules.
Ammonium, Deuterated
See Also
William M. Irvine
▶ Amino Acid
▶ Dinitrogen
▶ Mildly Reducing Atmosphere
Synonyms
▶ Nitrogen
▶ Nitrogen Fixation
Deuterated ammonium ion
▶ Titan
Definition
References and Further Reading The deuterated ammonium ion, NH3D+, has been
detected by radio astronomers toward a cold,
Altenhoff WJ, Batrla W, Huchtmeirs WK (1983) Radio dense ▶ molecular cloud core in the ▶ Milky
observations of Comet 1983 D. A & A 187:502
Way galaxy (Cernicharo et al. 2013;
Brandes JA, Hazen RM, Yoder HS (2008) Inorganic nitro-
gen reduction and stability under simulated hydrother- cf. ▶ Ammonia). Note that the symmetry of the
mal conditions. Astrobiology 8:1113–1126 corresponding principal ▶ isotopolog, NH4+,
88 Amoebae
leads to a zero electric dipole moment and hence See Also

no pure rotational transitions, making its inter-
stellar detection very difficult. Large isotopic ▶ Eukaryote
fractionation for deuterium/hydrogen isotopologs ▶ Protists
is expected, and indeed observed for many mol-
ecules, in cold molecular clouds.
Amorphous Carbon
See Also
Akira Kouchi
▶ Ammonia
Institute of Low Temperature Science, Hokkaido
▶ Isotopic Fractionation (Interstellar Medium)
University, Kita-ku, Sapporo, Hokkaido, Japan
▶ Molecules in Space
▶ Radio Astronomy
Keywords
Amorphous carbon; Carbon star; Carbonaceous
Cernicharo J, Tercero B, Fuente A, Domenech JL, chondrites; Cometary particles; Hydrogenated
Cueto M, Carrasco E, Herrero VJ, Tanarro I,
amorphous carbon; Interplanetary dust particles
Marcelino N, Roueff E (2013) Detection of the ammo-
nium ion in space. Astrophys J 771:L10–L13
Synonyms
Amoebae Glassy carbon; Vitreous carbon
Palaeobiogeology-Palaeobotany- Definition
Université de Liège, Liège, Belgium Amorphous carbon is a noncrystalline solid allo-
tropic form of carbon. There is no long-range
order in the positions of the carbon atoms, but
Definition some short-range order is observed. Chemical
bonds among atoms are a mixture of sp2- and
Amoebae are microscopic unicellular eukaryotes sp3-hybridized bonds with a high concentration
(▶ Protists) able to deform their cytoplasm to of dangling bonds. Because amorphous carbon is
move (amoeboid or crawling-like movement). thermodynamically in a metastable state and the
They represent a large diversity of unrelated ratio of sp2- and sp3-hybridized bonds is variable,
groups of eukaryotes. Some are surrounded by a the properties of amorphous carbon vary greatly
cell coat (glycocalyx); others are naked. Some are depending on the formation methods and condi-
pathogens. Others produce a mineral test made of tions (Silva and Ravi 2003). Amorphous carbon
siliceous plates, an organic test, or an agglutinated is often abbreviated as “a-C.”
test made of external organic or mineral particles
(thecamoebae or testate amoebae). Some amoebae
demonstrate a social behavior when several indi- Overview
viduals join to form complex multicellular struc-
tures such as slugs or fruiting bodies. The oldest In the laboratory, amorphous carbon can be pro-
fossil amoeba reported so far is 750 Ma old. duced by physical vapor deposition, chemical
Amorphous Carbon 89
vapor deposition, sputtering, and ion irradiation processing (UV photons and cosmic rays) of icy
of diamond or graphite. The structure of amor- grains in interstellar molecular clouds
phous carbon has been analyzed by X-ray and (Greenberg 1998; Kouchi et al. 2005). Amor- A
electron diffraction methods. The ratio of sp2- phous carbon grains have also been found in the
and sp3-hybridized bonds can be determined by matrix of carbonaceous chondrites (Brearley
electron energy loss spectroscopy, X-ray 2008). These grains are essentially made of pure
photoelectron spectroscopy, and Raman spec- carbon embedded in an amorphous silicate
troscopy. Amorphous carbon whose dangling matrix. It has been proposed that these grains
bonds are terminated with hydrogen is called were originally primitive macromolecular
hydrogenated amorphous carbon (a-C:H). organic material that has undergone mild thermal
Depending on the sp2 and sp3 ratios, the proper- metamorphism in the parent bodies of carbona-
ties of amorphous carbon differ greatly. When a ceous chondrites.
significant fraction of sp3 bonds is present in
amorphous carbon, this is called tetrahedral
amorphous carbon (ta-C) or diamond-like car-
bon. Tetrahedral amorphous carbon is hard, See Also
transparent, and electrically insulating and has
higher density than a-C and a-C:H. ▶ Insoluble Organic Matter
In space, the occurrence of amorphous carbon ▶ Kerogen
is observed in circumstellar envelopes around ▶ Molecular Cloud
carbon stars. When carbon stars lose mass to ▶ Organic Refractory Matter
stellar winds, carbonaceous materials, such as ▶ Polycyclic Aromatic Hydrocarbon
polycyclic aromatic hydrocarbons (PAH), SiC,
and amorphous carbon (a-C/a-C:H), that con-
dense in their extended atmospheres are released
to the interstellar medium. The conditions for the References and Further Reading
formation of amorphous carbon (a-C) grain have
been investigated theoretically (Gail and Blanco A et al (1990) Amorphous carbon and carbona-
Sedlmayr 1984), and the occurrence of amor- ceous materials in space II.-Astrophysical implica-
tions. Nuovo Cimento C 13:241–247
phous carbon (a-C) and SiC has been deduced Brearley AJ (2008) Amorphous carbon-rich grains in the
by observing the spectra of carbon stars (Blanco matrices of the primitive carbonaceous chondrites,
et al. 1990). ALH77307 and Acfer 094. Lunar Planet Sci
Very recently, amorphous carbon has been XXXIX:1494
Gail H-P, Sedlmayr E (1984) Formation of crystalline and
found in various extraterrestrial materials. amorphous carbon grains. Astron Astrophys
Cometary particles from comet 81P/Wild 2, cap- 132:163–167
tured by NASA’s Stardust mission, were ana- Greenberg JM (1998) Making a comet nucleus. Astron
lyzed by transmission electron microscopy, and Astrophys 330:375–380
Kouchi A et al (2005) Novel routes for diamond formation
a small amount of amorphous carbon grains less in interstellar ices and meteoritic parent bodies.
than 200 nm in size was found (Matrajt Astrophys J 626:L129–L132
et al. 2008). In interplanetary dust Matrajt G et al (2008) Carbon investigation of two Star-
particles (IDPs), investigated with Raman and dust particles: a TEM, NanoSIMS, and XANES study.
Meteor Planet Sci 43:315–334
infrared spectroscopy, the dominant type of Muñoz Caro GM et al (2006) Nature and evolution of the
carbon is found to be either a form of amorphous dominant carbonaceous matter in interplanetary dust
carbon (a-C) or of hydrogenated amorphous particles: effects of irradiation and identification with a
carbon (a-C:H), depending on the type of type of amorphous carbon. Astron Astrophys
459:147–159
IDP (Muñoz Caro et al. 2006). It has been pro- Silva S, Ravi P (eds) (2003) Properties of amorphous
posed that amorphous carbon in cometary parti- carbon, institution of engineering and technology.
cles and IDPs was formed by energetic INSPEC, London
90 Amorphous Solid
temperatures (400–600 C) and pressures

Amorphous Solid (200–900 MPa). Rocks in most Archean gneiss
belts are metamorphosed at the amphibolite
William M. Irvine facies. The name-giving rock is amphibolite, a
University of Massachusetts, Amherst, MA, USA dense and dark green to black rock generated by
metamorphism under moderate temperature
(ca. 500 C) and pressure (1 GPa) from a mafic
Definition (basaltic) protolith or, more rarely, from impure
dolostone (carbonate rock). It consists mainly of
An amorphous solid lacks long-range order in the hornblende, a type of amphibole, with lesser pro-
positioning of its constituent atoms; glass is an portions of plagioclase, and in some cases biotite,
example. This contrasts with a crystalline solid, epidote, titanite, and iron oxides. Amphibolite is
where such order is present, e.g., quartz. Both the a common constituent of metamorphosed oceanic
ices and the silicates in ▶ interstellar dust grains crust or of mafic intrusions in orogenic belts.
are typically amorphous, although crystalline sili-
cates are present in some circumstellar and come-
tary dust. The conversion of amorphous to
See Also
crystalline water ice has often been invoked as an
energy source in cometary outbursts at large helio-
▶ Metamorphic Rock
centric distances. The presence of crystalline sili-
▶ Oceanic Crust
cates (presumably formed in the hot and dense
inner solar system, possibly under the action of
energetic particles from the young Sun) in
▶ comets, which are formed in the cold outer
part of the solar system, suggests that mixing of Amphiphile
material was important in the ▶ solar nebula.
David Deamer
Department of Chemistry, University of
California, Santa Cruz, Santa Cruz, CA, USA
See Also
▶ Comet
Synonyms
▶ Interstellar Ices
Detergent; Lipid; Surfactant
▶ Solar Nebula
Definition
Amphibolite Facies An amphiphile is a molecule having both a

hydrophobic nonpolar group and a hydrophilic
Nicholas Arndt polar group. The nonpolar hydrophobic portion
ISTerre, Université Grenoble Alpes, France of the molecule is typically a hydrocarbon chain
ranging from 10 to 20 or more carbon atoms in
length, and the polar moiety can be a carboxylic
Definition acid, phosphate, sulfate, amine, or alcohol group,
among other possibilities. Examples of
Amphibolite facies refers to rocks formed under amphiphiles are fatty acids, detergents, and
metamorphic conditions of moderate to high all lipids including phospholipids and sterols.
Amphoteric Compounds 91
All amphiphiles are surface active and form Amphiphilic molecules can assemble in vari-
monolayers at air-water interfaces. Some amphi- ous solvents. Phospholipids are typical amphi-
philes, particularly those with a single hydrocar- philic biomolecules, having hydrophilic heads A
bon chain, assemble into ▶ micelles in aqueous including phosphate ions or charged tertiary or
solutions. Other amphiphiles with two hydrocar- quaternary amines and hydrophobic tails includ-
bon chains, for instance, phospholipids, typically ing fatty acids. Many phospholipids form lipid
self-assemble into bilayer membranes that are the bilayer membranes in water, where the polar
permeability barriers defining most forms of cel- heads face toward the solvent and hydrophobic
lular life. Amphiphilic molecules resembling tails which aggregate to form the inner part of the
fatty acids are present in carbonaceous meteorites vesicle.
and are plausible membrane-forming compo-
nents of the first living cells.
See Also
See Also
▶ Hydrophobicity
▶ Membrane
▶ Lipid Bilayer
▶ Self-Assembly
▶ Self-Assembly
▶ Self-Assembly, Biological
Amphiphilicity Ampholytes
Kensei Kobayashi
▶ Amphoteric Compounds
Definition
Amphoteric Compounds
Amphiphilicity refers to the property of some
Kensei Kobayashi
molecules to have an affinity to two phases and
most notably in biochemical systems the affinity
to both a polar solvent phase (in this case water)
and hydrophobic phase (such as the interior of
cell membranes or proteins). Amphiphilic mole-
cules usually contain both hydrophobic (e.g., Synonyms
benzyl or alkyl) and hydrophilic groups (e.g.,
-OH, -NH2 and -COOH). Amphiphilic molecules Ampholytes
are often useful as surfactants. Sodium
dodecylbenzene sulfonate is a typical amphi-
philic molecule used as a laundry detergent or Definition
shampoo and complexes hydrophobic substances
such as dirt and oil with its hydrophobic An amphoteric compound is a compound that can
dodecylbenzene moiety (C12H25-C6H4-) and is act both as an acid and as a base. Some metal
dispersed by affinity of the sulfonate moiety oxides or metal hydroxides, such as aluminum
(-SO3H) to water. oxide (Al2O3), show amphotericity:
92 Amplification (Genetics)
With a base : Al2 O3 þ 2 NaOH þ 3H2 O

Amplification (Genetics)
Ð 2 Na AlðOHÞ4
Carlos Briones
With an acid : Al2 O3 þ 6 HCl Ð 2 AlCl3 þ 3 H2 O Centro de Astrobiologı́a (CSIC/INTA),
Consejo Superior de Investigaciones Cientı́ficas,
Some organic compounds also show Madrid, Spain
amphotericity. These include amino acids, com-
pounds having both carboxylic group(s) and
amino group(s). For instance, glycine is predom- Definition
inantly present as a zwitterion (+NH3-CH2-
COO) in circumneutral aqueous solution, and In molecular biology, amplification is a process
it can neutralize either an acid or a base as by which a ▶ nucleic acid molecule is enzymat-
follows: ically copied to generate a progeny population
with the same sequence as the parental one. The
most widely used amplification method is the
With a base:þ H3 NCH2 COO þ NaOH ▶ polymerase chain reaction (PCR). The result
Ð H2 NCH2 COO Naþ þ H2 O of a PCR amplification of a segment of ▶ DNA
is called an “amplicon.” Nucleic acids can also
be amplified in an isothermal reaction involving
With an acid:þ H3 NCH2 COO þ HCl a reverse transcriptase, which copies
▶ RNA!DNA, and a DNA-dependent RNA
Ð Clþ H3 NCH2 COOH
polymerase, which transcribes DNA!RNA. Iso-
thermal amplification does not generate double-
Amino acids and their polymers (proteins) stranded DNA, and it is mainly used for copying
dissolved in aqueous solution possess both RNA. Ligase-based methods, including the
positively and negatively charged groups. so-called ligase chain reaction (LCR), can be
In acidic solution, there are typically more also used for specific DNA or RNA amplification.
positively charged groups, while there are A fourth general method for nucleic acid ampli-
typically more negatively charged groups in fication involves ▶ cloning the selected DNA
basic solution, though this depends somewhat molecule into bacterial or eukaryotic cells,
on the sequence of the protein. At a defined allowing them to reproduce, and collecting the
pH called the isoelectric point (pI), amino amplified DNA.
acids or proteins have balanced positive and
negative charges. At the isoelectric point of a
protein, its hydrophobicity becomes
maximum, and its solubility to water becomes See Also
minimum.
▶ Cloning
▶ DNA
▶ Nucleic Acids
See Also ▶ Plasmid
▶ Polymerase Chain Reaction
▶ Amino acid ▶ Replication (Genetics)
▶ Zwitterion ▶ RNA
Anaerobe 93
Definition
Anabolism
Anaerobes are organisms that do not require oxy- A
Juli Peretó gen to obtain energy or to grow. Anaerobic
Institut Cavanilles de Biodiversitat i Biologia metabolism is restricted to microorganisms,
Evolutiva, Universitat de València, València, both prokaryotic (▶ Bacteria and ▶ Archaea)
Spain and eukaryotic (yeast, microsporidia), although
an anaerobic multicellular organism (phylum
Loricifera) has been recently discovered in
marine sediments.
Synonyms
Biosynthesis
Overview
There are two main categories of anaerobic

Definition microorganisms: (1) facultative anaerobes that
can use oxygen for ▶ respiration if it is
Anabolism is the subset of metabolic networks by present but in its absence obtain energy from
which cell components are derived from ▶ fermentation (such as enterobacteria or
organic or inorganic precursors. Anabolism yeasts), ▶ anaerobic respiration (some Pseudo-
requires a source of energy – usually in the form monas, Thiobacillus, Bacillus, and many others),
of ATP – and reducing power, usually as and anoxygenic photosynthesis (some
NADPH. Proteobacteria) and (2) obligate anaerobes,
which never use oxygen. These can, in turn, be
divided into two subcategories: (a) strict or
See Also obligate anaerobes, for whom oxygen is poison-
ous (i.e., oxygen is extremely toxic to
▶ Assimilative Metabolism ▶ methanogens), and (b) aeroduric or
▶ Catabolism aerotolerant anaerobes that can grow in the pres-
▶ Metabolism ence of oxygen, although they never use it (i.e.,
bacteria involved in lactic acid fermentations).
Cultivating strict anaerobes in the laboratory is
an arduous task due to their extreme sensitivity to
oxygen. Anaerobic jars or chambers are neces-
Anaerobe
sary for the isolation and growth of methanogenic
archaea, sulfur-reducing bacteria, or bacteroides.
José Luis Sanz
Some anaerobes are etiological agents of
Departamento de Biologı́a Molecular,
important diseases, such as tetanus (Clostridium
Universidad Autónoma de Madrid,
tetani), botulism (Clostridium botulinum), chol-
Madrid, Spain
era (Vibrio cholera), salmonellosis and typhoid
fever (Salmonella enterica), or peptic ulcers
(Helicobacter pylori). Others, such as the lactic
Synonyms acid fermenters Lactobacillus and Lactococcus,
are involved in the production of food from dairy
Non-aerobic (yogurt, cheese, kefir, sour cream), vegetables
94 Anaerobic Photosynthesis
(sauerkraut, olives, pickles), or meat (sausages).

Some yeast (Saccharomyces) are responsible for Anaerobic Respiration
bread, beer, and wine production. Finally, the
methanogenic archaea carry out the last step of Juli Peretó
anaerobic degradation of organic matter in the Institut Cavanilles de Biodiversitat i Biologia
absence of oxygen and, therefore, play a key Evolutiva, Universitat de València,
role in anaerobic wastewater treatment and València, Spain
biomethanization of municipal solid waste pro-
cesses. It is important to underline that Earth’s
atmospheric O2 is of biological origin, and for an Definition
extended period of biological evolution, includ-
ing the period in which the ▶ origin of life is Anaerobic ▶ respiration is a metabolic process in
suggested, the Earth remained strictly which oxidized organic compounds, such as
anaerobic. Anaerobes are of astrobiological inter- fumarate, or inorganic molecules, such as nitrate,
est because anaerobic conditions prevail on many sulfate, or ferric ion, serve as the terminal ▶ elec-
planets, for instance, ▶ Mars. tron acceptor of an electron transport chain.
See Also See Also
▶ Anaerobic Respiration ▶ Aerobic Respiration

▶ Anoxygenic Photosynthesis ▶ Electron Acceptor
▶ Archaea ▶ Respiration
▶ Bacteria
▶ Fermentation
▶ Mars
▶ Methanogens Analog Sites
▶ Origin of life
▶ Respiration ▶ Terrestrial Analog
Madigan M, Martinko J, Dunlap P, Clark D (2009) Brock

biology of microorganisms, 12th edn. Person Education, Angular Diameter
Benjamin Cummings, San Francisco, Chapters 18, 21
Sowers KR, Noll KM (1995) Techniques for anaerobic
growth. In: Robb FT, Place AR, Sowers KR, Schreier Daniel Rouan
HJ, Dassarma S, Flischmann EM (eds) A laboratory LESIA, Observatoire Paris-Site de Meudon,
manual: methanogens. Cold Spring Harbor Laboratory Meudon, France
Press, New York, pp 15–47
Willey JM, Sherwood LM, Woolverton CJ (2008) Pres-
cott, Harley, and Kleins. Microbiology, 7th edn.
McGraw-Hill, Boston, Chap. 9
Definition
The angular diameter of a celestial object, seen

from Earth, is the apparent diameter measured in
Anaerobic Photosynthesis angular units. Planets in the solar system have
typical angular diameters between a few arcsec
▶ Anoxygenic Photosynthesis up to 50 arcsec (0.25 m-radians).
Angular Momentum 95
References and Further Reading a and b in the direction given by the right-
hand rule.
Arrhenius S (1903) Die Verbreitung des Lebens im Variations of velocity are produced by forces,
Weltenraum. Umschau 7:481–485 A
in accordance with Newton’s second law of
Arrhenius S (1908) Worlds in the making: the evolution of
the universe. Harper & Row, New York dynamics. If forces that apply on the point are
aligned with the position vector r, they are called
central, the system is invariant under rotation,
and its angular momentum is conserved. As
Angular Momentum ▶ gravitation is a central force, this conservation
occurs frequently in astronomy.
Jérôme Perez When the system is extended, such as a solid
Applied Mathematics Laboratory, ENSTA planet described by a distribution of points whose
ParisTech, Paris Cedex 15, France relative distances are fixed, the total angular
momentum is the sum of the contributions of all
these points. In this case, it is distributed between
Keywords the spin of the planet itself and the angular
momentum of its orbit.
Rotating body The conservation of angular momentum of
celestial bodies is a fundamental tool for analyz-
ing their properties. For example:
Definition
• In a two-body problem (see ▶ Gravitation),
if one of the two bodies is much heavier than
In mechanics, angular momentum is the vector
the other, the conservation of angular momen-
cross product between the position vector and the
tum implies Kepler’s third law (see ▶ Orbital
momentum vector of a point mass system. This
Resonance) and allows us to obtain the value
definition can be extended to a solid by
of the large mass from observations of the
summation.
period and of the semimajor axis of the
small mass.
• If a planet is found to rotate slower than
Overview expected, one can suspect that this planet
is accompanied by a satellite, because the
The movement of a point mass m is defined by its total angular momentum is shared between
position r and its velocity v. These quantities are the planet and its satellite in order to be
vectors relative to some reference system. This conserved.
movement splits into two parts: a movement of • The tidal torque the Moon exerts on the Earth
translation and a movement of rotation. The implies a slowing down of the rotation rate of
amount of movement is measured by the linear the Earth (at about 42 ns/day). As a conse-
momentum (impulsion) p = mv (for simple quence and because the total angular momen-
cases), which is a conserved quantity for a trans- tum of the whole system is conserved, the
lation invariant system. The amount of rotation is distance between the Earth and Moon gradu-
measured by the angular momentum L = r p, ally increases by 4.5 cm/year.
which is a conserved quantity for a rotation
invariant system. Note that the vector cross prod-
uct a b ¼ ab sin y n , where y is the smaller See Also
angle between a and b (0 y 180 ), a and
b are the magnitudes of vectors a and b, and n is a ▶ Gravitation
unit vector perpendicular to the plane containing ▶ Orbital Resonance
96 Animalcules
See Also
Animalcules
▶ Circumstellar Chemistry
Stéphane Tirard ▶ Molecules in Space
Centre François Viète d’Histoire des ▶ Photochemistry
Sciences et des Techniques EA 1161, Faculté
des Sciences et des Techniques de Nantes,
Nantes, France
Annefrank
Definition
Stefano Mottola
German Aerospace Center (DLR), Institute of
From the seventeenth century to the early nine-
teenth century, animalcule meant the very small
living beings that were observed through a micro-
scope. The famous microscopist, Antony van
Definition
Leeuwenhoek (1632–1723), one of the major
improvers of this instrument during the second
(5535) Annefrank is a small S-class, main-belt
part of the seventeenth century, used the expres-
asteroid named after the Jewish victim of Nazi
sion spermatic animalcules.
persecution famous for her diary. The Discovery
spacecraft Stardust encountered the asteroid on
November 2, 2002 on its route to comet Wild
See Also
2. During the fast flyby, the space probe recorded
an image sequence lasting about 15 min and
▶ Bacteria
consisting of over 70 images. Although the imagery
▶ Protists
had a comparatively low resolution in the range of
300–185 km/pixel and covered only about 40 % of
the surface, the sequence revealed a body with
approximate dimensions of 6.6 5.0 3.4 km
Anion and an angular appearance, reminiscent of a contact
binary or of a re-accumulated pile of fragments.
Steven B. Charnley
Anorthosite
Nicholas Arndt
Definition ISTerre, Université Grenoble Alpes, France
An anion is an atom or molecule that has gained

an electron (e.g., CN, OH, C4H). Neutral Definition
molecules with large electron affinities (EAs)
can attach an electron in a variety of chemical Anorthosite is a magmatic intrusive rock. It is
reactions, such as electron photo attachment. light colored (leucocratic) and has a medium to
Long carbon-chain molecules have large EA coarse grain size (phaneritic). It is mainly com-
values, and C6H was the first anion discovered posed of plagioclase (andesine, labradorite,
in the interstellar medium in 2006. bytownite) and minor pyroxene, olivine, and
Antarctic Continent 97
iron-titanium oxides (ilmenite, magnetite). Prote-

rozoic anorthosite forms large massifs associated Anoxygenic Photosynthesis
with granitoids (North America, Scandinavia). A
Archean coarse-grained (megacrystic) anortho- Juli Peretó
site occurs in intrusions (dikes and sills) and Institut Cavanilles de Biodiversitat i Biologia
flows of basaltic composition. Evolutiva, Universitat de València, València,
Anorthosite is a common constituent of the Spain
lighter surfaces of the Moon called lunar high-
lands or terrae. Formation of anorthosite requires
the concentration of plagioclase from mafic
magma by flotation in a magma ocean (as is pro- Synonyms
posed to have occurred on the Moon), ascent of
plagioclase-rich mushes, or low-pressure crystal- Anaerobic photosynthesis
lization in magma chambers.
See Also Definition
▶ KREEP Anoxygenic ▶ photosynthesis is a bacterial

▶ Mafic and Felsic photosynthesis that occurs under anaerobic
▶ Moon, The conditions, using the photosynthetic electron
transport chain in a noncyclic mode and
reduced inorganic electron donors, such as
hydrogen sulfide, hydrogen, or ferrous ion, as
▶ electron donors. There are also cases of
Anoxic anaerobic photosynthetic electron transport
chains acting cyclically; in this case, the
Ricardo Amils
generation of reducing power is not needed or
Departamento de Biologia Molecular,
it is decoupled from the photosynthetic
Universidad Autónoma de Madrid,
reaction. The prototypical noncyclic
Madrid, Spain
anoxygenic photosynthesis is present in green
bacteria.
Definition
Anoxic is a term used to describe a condition,

See Also
environment, or habitat depleted of oxygen.
▶ Electron Donor
▶ Photosynthesis
See Also
▶ Photosynthesis, Oxygenic
▶ Anaerobe
Anoxic Ocean Antarctic Continent
▶ Sulfidic Oceans ▶ Antarctica

98 Antarctica
ice-free Antarctica McMurdo Dry Valleys reflect

Antarctica surface conditions similar to those of the Mars
surface. Dry Valleys are located mainly in the
Daniele L. Pinti Victoria Land west of McMurdo Sound. Dry val-
GEOTOP Research Center for Geochemistry and leys are so named because of their extremely low
Geodynamics, Université du Québec à Montréal, humidity and their lack of snow and ice cover.
Montréal, QC, Canada Their surfaces are covered by loose gravel and
show ice-wedge polygonal-patterned ground.
Polygonal-patterned ground is a geometric land-
Keywords form with characteristic honeycomb patterns
surrounded by ice crests that develop in periglacial
Glaciation; Ice sheets; Mars natural analog; regions which experience intense freezing and
Weathering thawing cycles. High Resolution Imaging Science
Experiment (HiRISE) camera on NASA’s ▶ Mars
Synonyms Reconnaissance Orbiter have clearly shown the
same patterns on high latitudes of Mars and at
Antarctic continent the bottom of some Mars crater. Antarctica Dry
Valleys and Mars mid-latitudes soil formation his-
Definition tories shares similarities, particularly involving
slow processes of sublimation and poleward
Antarctica is the ice-covered continent located in migration of water (Wentworth et al., 2005).
the Southern Hemisphere of the Earth. It is situ- Antartica permafrost hard surfaces are
ated almost entirely south of the Antarctic Circle privileged site of several Mars analog missions,
and is surrounded by the Southern Ocean. because the soils share physical similarities with
that of Mars. One of the most recent
Overview missions took place in the Marambio Island,
Western Antarctic Peninsula, where field explo-
Antarctica, the coldest and driest continent on ration, sample collection, instrument deploy-
Earth, is technically considered a desert, with ment, and spacesuit testing were carried out on
only 200 mm per year of average precipitation. a Mars-like rocky, permafrost-rich landscape
Antarctica also recorded the coldest temperature (Rask et al., 2012).
on Earth measured so far, of 89.2 C at Vostok
station. Ninety-eight percent of Antarctica’s
14 million km2 surface area is covered by a See Also
1.6 km thick ice sheet, corresponding to 90 % of
the world’s ice. It has been covered by ice for the ▶ Europa
past 15 Ma. About 400 subglacial lakes lie at the ▶ Mars Analogue sites
base of the continental ice sheet, the best known ▶ Vostok, Subglacial Lake
being Lake Vostok. Lake Vostok has remained
isolated for 14 million years, making it a valuable
analog for exploring deep biosphere niches. References and Further Reading
Antarctica ice surface shares similarities with
Rask JC, De León P, Marinova MM, McKay CP
those of ▶ Jupiter’s moon ▶ Europa and ▶ Mars. (2012) The exploration of Marambio Antarctica as a
Antarctica has been for long time a privileged Mars analog. 43rd Lunar and Planetary Science Con-
natural analog terrain for studying morphological ference Abstract, 2455
Wentworth SJ, Gibson EK, Velbel MA, McKay DS
processes shaping the Mars surface or for testing
(2005) Antarctic Dry Valleys and indigenous
instrumentation and field exploration procedures weathering in Mars meteorites: implications for
for future manned Mars missions. Particularly, water and life on Mars. Icarus 174:383–395
Anticodon 99
Antibiotic Antibody
A
Ricardo Amils Juli Peretó
Departamento de Biologia Molecular, Institut Cavanilles de Biodiversitat i Biologia
Universidad Autónoma de Madrid, Evolutiva, Universitat de València, València,
Madrid, Spain Spain
Definition
Synonyms
Antibody is a complex ▶ protein
Antimicrobial agent; Functional inhibitor (immunoglobulin) produced as a response to a
chemical agent (antigen) as a part of a defensive
system (immune system) in multicellular ani-
mals. The combination of antibody and antigen
Definition is specific (albeit not necessarily absolute),
non-covalent, and reversible. There are many
Antibiotics are chemical substances produced methodological applications of antibodies,
by a wide range of microorganisms, among either as a heterogeneous population of immuno-
them fungi and bacteria, that kill or globulins (polyclonal antibodies) or as a homo-
inhibit the growth of other organisms. A large geneous preparation (monoclonal antibodies).
number of antibiotics have been identified in Antibodies show a broad applicability in biotech-
nature, most of them as products of secondary nology, including the development of affinity
metabolism. Antibiotic producers must be biosensors.
resistant to the active form of the
antibiotic. Important targets of antibiotics are
the synthesis of ▶ cell membrane and
▶ cell wall, replication, ▶ transcription, and See Also
▶ translation. Antibiotics are considered
regulators of microbial populations rather ▶ Biosensor
than part of microbial warfare. The susceptibility ▶ Protein
of organisms to individual antibiotics or
other chemotherapeutic agents varies signifi-
cantly and is the base of their
pharmacological use.
Anticodon
Juli Peretó
Institut Cavanilles de Biodiversitat i Biologia
See Also
Evolutiva, Universitat de València,
València, Spain
▶ Cell Membrane
▶ Cell Wall
▶ Replication (Genetics)
▶ Ribosome Definition
▶ Sporulation
▶ Transcription Anticodon is a triplet of nucleotides in a tRNA,
▶ Translation complementary to a codon in the mRNA.
100 Antimicrobial Agent
See Also
Apex Basalt, Australia
▶ Codon
▶ Genetic Code Nicholas Arndt
▶ RNA ISTerre, Université Grenoble Alpes, France
▶ Translation
▶ Wobble Hypothesis (Genetics)
Definition
The Apex Basalt is a ca. 3.46-Ga-old formation

comprising tholeiitic pillow basalts, komatiitic
Antimicrobial Agent basalts, and komatiites intercalated with thin
chert layers. It is located near Marble Bar in the
▶ Antibiotic ▶ Pilbara Craton of Western Australia. ▶ Micro-
fossils, morphological biomarkers, and filamen-
tous carbon structures in the lower chert beds
have been interpreted as fossil prokaryotes
(mainly cyanobacteria but also thermophiles)
AOGCM and are claimed to represent the oldest fossil
record of life on Earth. For this reason, outcrops
John Lee Grenfell of this formation are considered one of the most
German Aerospace Center (DLR), Berlin, important astrobiological sites on Earth.
Germany
See Also
Synonyms ▶ Apex Chert
▶ Apex Chert, Microfossils
Atmosphere-ocean general circulation model ▶ Mars Analogue Sites
▶ Microfossils
▶ Pilbara Craton
Definition
An AOGCM refers to a 3D numerical model

which solves the central conservation equations, Apex Chert
e.g., mass, momentum, and energy, to derive the
characteristic global-scale fluid dynamical flow Tanja Elsa Zegers
(the “general circulation”) as well as temperature Paleomagnetic Laboratory, Institute of Earth
of a planetary atmosphere and ocean. The atmo- Sciences, Utrecht University, Utrecht, CD,
sphere and ocean modules are coupled via surface The Netherlands
exchange fluxes of energy (e.g., via evaporation
and condensation) and momentum (e.g., via wind
stresses at the ocean surface). Definition
The 3.465 Ga Apex Chert is a chert unit within

the Apex Basalt in the Warrawoona Group,
See Also which is part of the oldest greenstone sequence
in the Pilbara granite-greenstone terrain. The
▶ GCM Apex Basalt is stratigraphically below the
Apex Chert, Microfossils 101
Strelley Pool Chert, a unit known for hosting the Definition

oldest ▶ stromatolite on Earth. In the Apex Chert,
small carbonaceous filaments with d13C as low as The Apex Chert is a bedded, microcrystalline A
22.5 to 25‰ were reported to represent evi- silica (SiO2) deposit interlayered with ▶ pillow
dence for ▶ cyanobacteria able to recycle inor- lavas and massive flows of the Apex Basalt For-
ganic carbon through ▶ RubisCO. The Apex mation, ▶ Pilbara Craton, Western Australia. The
Chert microfossils occur in rounded grains of micro- ▶ basalts were dated at 3,465–3,458 Ma. The
crystalline silica, which have been interpreted as origin of the chert is disputed, and interpretations
clasts in a conglomerate deposited in a wave-washed of primary silica deposition on the ocean floor or
beach or a stream mouth, an ideal environment for alternatively secondary, hydrothermal silicifica-
cyanobacteria. Subsequent work suggested that the tion (chertification) of clastic or carbonate sedi-
chert was deposited from hydrothermal fluids with a mentary and volcano-sedimentary rocks rival one
temperature higher than 250 C and that the another. The putative ▶ microfossils of Apex
microtextures may result from abiotic processes Chert are carbonaceous filaments found in
under those temperatures. If biogenic, microfossils ca. 3,465 Ma old chert lenses at the so-called
could represent remains of thermophile Schopf locality, Chinaman Creek near Marble Bar.
chemothrophs living close to hydrothermal vents.
Overview
See Also
The name of the famous “Schopf locality” where
▶ Apex Basalt, Australia the microfossils were found, derives from the
▶ Apex Chert, Microfossils American paleontologist and paleobiologist,
▶ Archean Traces of Life J. William (Bill) Schopf, who, at this site,
▶ Biomarker, Isotopic reported 11 morphological taxa of prokaryotic,
▶ Biomarkers, Morphological filamentous, and coccoidal microfossils embed-
▶ Carbon Isotopes as a Geochemical Tracer ded in chert clasts. At that time, the host rock was
▶ Cyanobacteria thought to constitute a sedimentary layer and
▶ Pilbara Craton later reinterpreted as sedimentary fill of a hydro-
▶ Rubisco thermal vein. The kerogenous (carbonaceous) fil-
▶ Stromatolites aments, up to several tens of micrometers long
and 1–20 mm wide, show in most cases a typical
cyanobacteria-like septation and terminal cells of
varying morphology (Fig. 1). They form single
Apex Chert, Microfossils cell chains and single coccoids that were
interpreted as the Earth’s oldest microfossils
Daniele L. Pinti1 and Wladyslaw Altermann2 (Schopf 1993). The morphology of the filaments
1
GEOTOP Research Center for Geochemistry and their organic carbon isotopic composition
and Geodynamics, Université du Québec à (d13C) ranging from 22 ‰ to 26 ‰ (whole
Montréal, Montréal, QC, Canada rock measurements), and the sedimentary envi-
2
Department of Geology, University of Pretoria, ronment interpreted as shallow marine, strongly
Pretoria, South Africa suggested that some of these filaments were
cyanobacteria. The hypothesis of cyanobacterial
life 3.5 billion years ago implied, for uniformi-
Keywords tarianism, that oxygenic photosynthesis might
have acted very early in the Earth’s history and
Apex Chert; Apex Basalt; Biomarkers; that life was already well advanced one billion
Cyanobacteria; Microfossils years after the Earth formation. Though the
102 Apex Chert, Microfossils
Apex Chert, Microfossils, Fig. 1 Microfossils from the drawings. All at a magnification as shown in (A). (a–e)
early Archean Apex Chert of Australia (From Schopf Primaevifilum amoenum; (f–j) Primaevifilum
1993). Microfossils (A-J, holotypes) with interpretative conicoterminatum (with conical terminal cells)
evidence of microfossils in Apex Chert, as well as Geology of the Schopf Locality

the paleoenvironmental and depositional condi- The Apex Chert is a unit within the 4 km thick
tions, have been later highly debated, the Apex Apex Basalt Formation of the Salgash Subgroup
Chert and its putative microfossils have greatly of the Warrawoona Group of ▶ Pilbara Craton,
contributed to a general interest in the origin Western Australia. The Apex Basalt consists of
and evolution of life and in many ways toward ▶ greenschist facies metamorphosed ▶ basalts,
astrobiology, encouraging the search for extrater- komatiitic basalts, and serpentinized ▶ perido-
restrial life, the improvement of our understand- tites and minor felsic volcaniclastic rocks with
ing of abiotic and biotic, evolutionary and locally intruded dolerite sills which form part of
taphonomic (processes by which organisms the Marble Bar ▶ greenstone belt.
become fossilized) processes, and to the develop- The “Schopf locality” (Schopf and Packer
ment of careful investigation methods and defi- 1987; Schopf 1993), north of Marble Bar, is
nition of unambiguous biosignatures. well known for its chert formations. The Apex
Chert, with an assigned age of 3,465 5 Ma, is a division that morphologically resembled
bedded unit consisting of up to 10 m thickness of cyanobacteria (Fig. 1). Next to the filaments,
white, gray, and black-layered chert, interbedded hundreds of solitary unicell-like spheroidal struc- A
with felsic tuff, which contains sills of massive tures resembling coccoidal microfossils were
black silica. The bedded deposits overlie a swarm identified. Based on morphology, Schopf and
of weakly radiating black silica veins that extend Packer (1987) and Schopf (1992, 1993, 2006)
up to 750 m stratigraphically down into recognized 11 morphotypes of putative microfos-
metabasalts, but which do not penetrate above sils in the Apex Chert, as listed below:
the bedded chert horizon, cut by an unconformity.
The veins themselves are composed of several 1. Narrow unbranched septate prokaryotic fila-
phases of intrusive silica that vary in color ments incertae sedis cf. bacteria?
from very dark blue-black, through shades of (Archaeotrichion septatum)
blue-gray, to white. They migrate sidewards 2. Narrow unbranched septate prokaryotic fila-
into the sedimentary layers replacing them ments incertae sedis cf. bacteria?
with silica. The veins comprise dominantly (Eoleptonema apex)
massive dark blue-black silica, but can include 3. Narrow unbranched septate prokaryotic fila-
multiple generations of dark gray to black ments incertae sedis cf. bacteria? or
silica to white quartz, core zones of felsic cyanobacteria? (Primaevifilum minutum)
tuff ▶ breccia, and phreatomagmatic 4. Narrow unbranched septate prokaryotic fila-
breccias with a jigsaw puzzle fit with exploded ments incertae sedis cf. bacteria? or
fragments at their tops (Van Kranendonk and cyanobacteria? (Primaevifilum delicatulum)
Pirajno 2004). The bedded chert of the Schopf 5. Intermediate-diameter unbranched septate
microfossil locality is the stratigraphically lowest prokaryotic filaments incertae sedis
of five bedded chert units within the pillowed cf. cyanobacteria? (Primaevifilum amoenum)
Apex Basalt. 6. Intermediate-diameter unbranched septate
prokaryotic filaments having disk-shaped
Microfossils at the Schopf Locality medial cells incertae sedis
The “Schopf locality,” where the microfossils cf. cyanobacteria? (Archaeoscillatoriopsis
were discovered, is controversially within a disciformis)
black chert vein radiating from the bedded chert 7. Broad unbranched septate prokaryotic fila-
unit and is not from the bedded Apex Chert Unit ments having conical end cells incertae
itself. However, the fossiliferous specimen sedis cf. cyanobacteria? (Primaevifilum
deposited by Schopf at the Natural History conicoterminatum)
Museum, London, shows bedded structure and 8. Broad unbranched septate prokaryotic fila-
brownish color, implying that they come from ments having equant medial cells incertae
the bedded part of the section. Microfossils sedis cf. cyanobacteria? (Primaevifilum
were discovered in rounded grains of microcrys- laticellulosum)
talline silica, apparently within one of the blue- 9. Broad unbranched septate prokaryotic fila-
black veins beneath the lowermost of the bedded ments incertae sedis cf. cyanobacteria?
chert units of the Apex Basalt Formation. Schopf (Archaeoscillatoriopsis grandis)
(1993, 1999) interpreted the grains as clasts of a 10. Broad unbranched markedly tapering septate
conglomerate deposited in a wave-washed beach prokaryotic filaments incertae sedis
or a stream mouth, an ideal environment for cf. cyanobacteria? (Primaevifilum
cyanobacteria. Fragments of stromatolites pro- attenuatum)
vide evidence that at least part of the clasts is of 11. Broad unbranched septate prokaryotic fila-
sedimentary origin. Schopf (1993) observed hun- ments having hemispheroidal end cells
dreds of filaments, tens of micrometers long and incertae sedis cf. cyanobacteria?
1–20 mm wide, some of them showing a septate (Archaeoscillatoriopsis maxima)
The Debate hardly have survived (Brasier et al. 2002), except

Remapping of the Marble Bar area including the for chemoautolithotroph thermophiles (Brasier
Schopf locality and detailed petrology and min- et al. 2006). Alternatively, an acid-sulfate
eralogy of the Apex Basalt Formation and Apex epithermal environment of alteration, syn- or
Chert Unit (Brasier et al. 2002, 2005; Van post-genetic with the precipitation of the chert,
Kranendonk and Pirajno 2004) revealed that the has also been suggested (van Kranendonk and
Apex Chert is largely a breccia infilling one of Pirajno 2004). The abundance of sulfate and
multiple generations of metalliferous hydrother- lack of argilitic alteration indicated depositional
mal veins. These veins crosscut pillow basalts temperatures up to 350 C. Recently, Pinti
and feed into, and are continuous with the over- et al. (2009) showed that medium-low tempera-
lying stratiform-bedded chert unit of the Apex ture weathering processes could explain the min-
Basalt Formation. eralogy of the Apex Chert so that high-
The discussion on the reality of Schopf’s temperature hydrothermal fluid-rock interactions
(1993) findings was triggered by claims that the are not required. Alternatively, the metal rich
filaments are branching, unlike prokaryotic fila- fluids might have passed at a much later time
ments, and do not contain carbon. Simulta- through these rocks.
neously, it was claimed that life did not exist on These observations invigorated the debate on
Earth prior to ca. 2,500 Ma (Brasier et al. 2002, the ▶ biogenicity of the carbonaceous filaments
2004). At reexamination, the Apex Chert, how- and their putative inclusion in the phylum
ever, was found to contain cellular-preserved cyanobacteria. Laser-Raman imagery of carbona-
kerogenous microfossil remains, revealing ceous filaments (Schopf et al. 2002, 2007; Brasier
advanced biostratonomic to metamorphic, tapho- et al. 2002) and disseminated carbonaceous
nomic changes. It was suggested that thermal (kerogenous) matter in the Apex Chert
alteration is the cause of taphonomic changes in (De Gregorio and Sharp 2006) gave controversial
cyanobacterial microfossils, resulting in the pre- results. Schopf et al. (2002) interpreted the car-
sent form of microfossil preservation in the Apex bon as of biological origin. Brasier et al. (2002)
Chert (Kazmierczak and Kremer 2002). The pre- proposed that it rather could be amorphous car-
served morphological variation indicates biolog- bon reorganized in the form of filamentous
ical behavior and fulfills the requirements for strains after devitrification processes of the chert
microfossil recognition (Buick 1990). Claims of veins. De Gregorio and Sharp (2006) suggested
branching of the filaments or of incomplete, that the carbonaceous material is similar in struc-
selective photomontages of the microstructures, ture to microfossil kerogen, but may also be pro-
mimicking a biological appearance (Brasier duced abiotically via Fischer-Tropsch-type
et al. 2002, 2004), result from misinterpretation (FTT) synthesis reactions, in an ancient hydro-
of auto-montages of photographs taken at differ- thermal vent. However, it has never been con-
ent depth of focus and superimposed on each firmed that the FTT process can produce
other (Fig. 2). However, the lack of assessment particulate carbon. Three-dimensional confocal
of the geological context of the Apex microfossil laser microscopy and Raman imagery demon-
assemblage together with the generally poor pres- strated that the structures are indeed cellular-
ervation due to possible biological, diagenetic, made filaments and coccoids (Schopf and
and metamorphic degradation cast some doubts Kudryavtsev 2005).
on the applied taxonomy in some cases The carbon isotopic composition is also contro-
(Altermann 2005). versial. The in situ, on single microfossils, mea-
Scanning electron microscopy (SEM) showed sured d13C values from 27 ‰ to 34 ‰ could
the presence of metals (Ni, Cu, Zn, Sn), sulfides, be related to photosynthesis (d13C = 25 ‰
barite, jarosite, alunite, phyllosilicates, and Fe 10 ‰; Schopf 2006), methanogenesis (Brasier
oxides, suggesting a high-temperature hydrother- et al. 2002), or abiotic FTT reactions (e.g.,
mal environment where microbial life could McCollom and Seewald 2006).
Apex Chert, Microfossils, Fig. 2 The same specimen of focus within the several tens of mm thick petrographic
Archaeoscillatoriopsis disciformis filament photographed thin section (from left to right and downwards). It
from the original material deposited by Schopf at the becomes clear that Schopf (1993) has shown only one
Natural History Museum, London. Upper left, as depicted filament located closer to the surface of the section,
by Schopf (1993), and upper right, as shown by Brasier while Brasier et al. (2002) have shown a sandwich photo-
et al. (2002): the difference created the impression that graph, including all depth of focus and exhibiting two
Schopf (1993) has manipulated the microphotographs in filaments coincidentally superimposed one above the
reality showing a branching and therefore impossibly other within the thickness of the section (Photograph by
cyanobacterial filament. The following lower micro- W. Altermann in M. in Brasier’s lab 2003)
graphs show the same filament at different depth of
Buick (1984) suggested that carbonaceous fil- Chert dyke or beds. Some clasts contain stromat-
aments in the silica swarm dykes of the North olitic laminae and relict carbonate minerals and
Pole and Marble Bar, including Apex Chert, were therefore must have been silicified during early
contaminants introduced in the microfracturing diagenesis at their source of origin (Fig. 3). The
of the silica veins during the tectonic uplift of the hydrothermal chert distinctly differs from these
region, 2.75 Ga ago. Pinti et al. (2009) observed clasts. However, in some places, hydrothermal
branched microstructures suggesting post- recrystallization strongly affects the clasts and
depositional colonization of microcracks and fis- they become almost non-discernible from the
sures by microbes. However, Schopf’s hydrothermal chert matrix. The clasts are thus
kerogenous microfossils are embedded in pri- clearly older than the hydrothermal dike
mary chert, in clasts deposited within the Apex (Altermann and Kazmierczak 2003; Altermann
Apex Chert, Microfossils, Fig. 3 Stromatolitic clast matter. The clast is cut by two parallel silica veinlets
within the microfossiliferus samples deposited by Schopf with pyrite enrichment (Photograph by W. Altermann in
at the Natural History Museum, London, exhibiting M. Brasier’s lab, 2003)
microbial lamination, pyrite grains, and dark organic
2007). The Apex Chert seems to have been Applications

affected by several hydrothermal and supergene
episodes of weathering, suggesting that it is Most of all the techniques developed for deter-
unlikely to have preserved any early forms of life. mining the biogenicity and syngenicity of
Nevertheless, it contains stromatolitic clasts, and Archean traces of life have been tested on Apex
stromatolites are known within this stratigraphic Chert (e.g., De Gregorio and Sharp 2006), and
succession. Moreover, even older microfossils several among them were specifically
and stromatolites were described from equally developed to resolve the dilemma of the Schopf
metamorphosed and altered shallow marine and microfossils (e.g., Schopf et al. 2002, 2005).
hydrothermal environments of the underlying This rock represents thus the best challenge
Dresser Formation (3,490 Ma) (Awramik for determining the reality of very ancient
et al. 1983; Ueno et al. 2001; Allwood et al. 2006). traces of life and developing successful method-
Whether the carbonaceous filaments of ologies and strategies of search for
J. William Schopf are genuine ancient fossilized extraterrestrial life.
prokaryotes (Schopf 1993; Altermann 2005),
later biological contamination (Pinti et al.
2009), or abiotic products (Brasier et al. 2005), Future Directions
this rock is still the most fascinating challenge in
Archean paleobiology and astrobiology. The critical The uniqueness of these microfossils constrains
point in views opposing the microfossil interpreta- the use of destructive methods for determining
tion is that most of these investigations were not the environmental context of deposition of this
made on the original specimen described by Schopf chert unit and the reality of these microfossils.
1993 and deposited in the Natural History Museum, New nondestructive techniques such as
London, but on rocks collected years later, in the NanoSIMS imagery of microfossils (Oehler
same outcrops but not exactly the same location. In et al. 2009) or a combination of analytical tech-
recent discussions, Schopf has reinforced his argu- niques could be useful for determining whether
ments for microfossils, introducing new data and the chemical structure of such putative microfos-
new investigation techniques (Schopf and sils is consistent with a biological origin
Kudryavtsev 2012; 2013, Pinti et al. 2013). (Derenne et al. 2008).
See Also Brasier M, Mcloughlin N, Green O, Wacey D (2006)

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Altermann W (2007) The early Earth’s record of enig- Schopf JW (1992) Paleobiology of the Archean. In:
matic cyanobacteria and supposed extremophilic bac- Schopf JW, Klein C (eds) The Proterozoic biosphere.
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cyanobacteria in extreme environments. Cellular ori- Schopf JW (1993) Microfossils of the early Archean apex
gin, life in extreme habitats and astrobiology (COLE) chert: new evidence of the antiquity of life. Science
11. Springer, Berlin, pp 759–778 260:640–646
Altermann W, Kazmierczak J (2003) Archean microfos- Schopf JW (1999) The cradle of life. Princeton University
sils: a reappraisal of early life on Earth. Res Microbiol Press, New York
154:611–617 Schopf WJ (2006) Fossil evidence of Archaean life. Phil
Awramik SM, Schopf JW, Walter MR (1983) Filamentous Trans R Soc B 361:869–885
fossil bacteria from the Archean of Western Australia. Schopf JW, Kudryavtsev AB (2005) Three-dimensional
Precambrian Res 20:357–374 Raman imagery of Precambrian microscopic organ-
Brasier MD, Green OR, Jephcoat AP, Kleppe AK, Van isms. Geobiology 3:1–12
Kranendonk MJ, Lindsay JF, Steele A, Grassineau NV Schopf JW, Kudryavtsev AB (2012) Biogenicity of
(2002) Questioning the evidence for Earth’s oldest Earth’s earliest fossils: a resolution of the controversy.
fossils. Nature 416:76–81 Gondwana Res 22:761–771
Brasier M, Green O, Lindsay J, Steele A (2004) Earth’s Schopf JW, Kudryavtsev AB (2005) Reply to the com-
oldest (similar to 3.5 Ga) fossils and the “Early Eden ments of D.L. Pinti, R. Mineau and V. Clement, and
hypothesis” questioning the evidence. Orig Life A.O. Marshall and C.P. Marshall on “Biogenicity of
Evol Biosph 34:257–269 Earth’s earliest fossils: a resolution of the controversy”
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Stoakes C (2005) Critical testing of Earth’s oldest Schopf JW, Kudryavtsev AB (2013) Reply to the com-
putative fossil assemblage from the 3.5 Ga Apex ments of D.L. Pinti, R. Mineau and V. Clement, and
chert, Chinaman Creek, Western Australia. Precam- A.O. Marshall and C.P. Marshall on “Biogenicity of
brian Res 140:55–102
108 Aphelion
Earth’s earliest fossils: a resolution of the contro-

versy”. Gondwana Res 23:1656–1658 Apolar Molecule
Schopf JW, Packer BM (1987) Early Archean (3.3- billion
to 3.5-billion-year-old) microfossils from
Warrawoona Group, Australia. Science 237:70–73 William M. Irvine
Schopf JW, Kudryavtsev AB, Agresti DG, Wdowiak TJ, University of Massachusetts, Amherst, MA, USA
Czaja AD (2002) Laser-Raman imagery of Earth’s
earliest fossils. Nature 416:73–76
Schopf JW, Kudryavtsev AB, Agresti DG, Czaja AD,
Wdowiak TJ (2005) Raman imagery: a new approach
Synonyms
to assess the geochemical maturity and biogenicity of
permineralized Precambrian fossils. Astrobiology
5:333–371 Nonpolar molecule
Schopf JW, Kudryavtsev AB, Czaja AD, Tripathi AB
(2007) Evidence of Archean life: stromatolites and
microfossils. Precambrian Res 158:141–155
Ueno Y, Maruyama S, Isozaki Y, Yurimoto H (2001) Definition
Early Archean (ca. 3.5 Ga) microfossils and 13C-
-depleted carbonaceous matter in the North Pole area, In interstellar chemistry, apolar molecules are
Western Australia. In: Nakashima S, Maruyama S,
molecules lacking a permanent electric dipole
Brack A, Windley BF (eds) Field occurrence
and geochemistry, in geochemistry and the moment. The lack of a dipole moment results
origin of life. Universal Academic Press, Tokyo, from the symmetry of the charge density distri-
pp 203–236 bution in the molecule. Such molecules have no
Van Kranendonk MJ, Pirajno F (2004) Geochemistry of
pure rotational transitions; hence, in the gas
metabasalts and hydrothermal alteration zones associ-
ated with c. 3.45 Ga chert and barite deposits. Impli- phase, they must be observed via their vibrational
cations for the geological setting of the Warrawoona or electronic transitions.
Group, Pilbara Craton, Australia. Geochem Explor
Environ Anal 4:253–278
See Also
▶ Polar Molecule
Aphelion
Daniel Rouan
LESIA, Observatoire Paris-Site de Meudon, Apollo Asteroid
Meudon, France
Alan W. Harris
DLR, Institute of Planetary Research, Berlin,
Definition Germany
The aphelion is the point on a body’s orbit around

the Sun (planets, comets, asteroids) where the
Definition
body is farthest from the Sun.
An Apollo ▶ asteroid is a near-Earth asteroid
with a semimajor axis of more than 1 astronomi-
See Also cal unit (AU) and a perihelion distance of less
than 1.017 AU (the Earth’s aphelion distance).
▶ Keplerian Orbits The ▶ orbit of such an ▶ asteroid may intersect
▶ Orbit that of the Earth, giving rise to an impact hazard.
▶ Periastron Apollo asteroids are named after the asteroid
Apollo Mission 109
1862 Apollo, which is the first to be discovered exploration with President J. F. Kennedy’s
having these dynamical characteristics. announcement in 1961 of sending an American
safely to the Moon before the end of the decade A
and at same time the progress in the technical
See Also capabilities of space transportation. The Apollo
program ultimately placed 12 men on the lunar
▶ Asteroid surface. In 1972, with Apollo 16 and 17, the era of
▶ Near-Earth Objects human exploration beyond Earth’s orbit was ter-
▶ Orbit minated and so far it has not been resumed.
Overview
Apollo Mission
The Apollo missions to the Moon were
performed between 1968 and 1972 (Table 1).
Gerda Horneck
The Apollo missions were the first and so far
DLR German Aerospace Center, Institute of
only human space missions beyond Earth’s orbit.
Aerospace Medicine, Radiation Biology,
They provided in-depth knowledge of the geol-
Köln, Germany
ogy of the Moon (Schaber 2005) and biomedical
data on human health issues during space flight
(Johnston et al. 1975). Biological responses to the
Keywords
parameters of outer space were studied in the
following experiments:
Biological effects of space; Exposure experi-
ments; Human space flight; Lunar missions
• ▶ Biostack experiments on board of the
Apollo 16 and 17 Command Module on the
responses of a variety of biological systems in
Synonyms
resting state to the heavy ion component of
cosmic rays (B€ucker and Horneck 1975)
NASA lunar landing mission
• ALFMED experiment during the Apollo
16 and 17 mission that demonstrated that the
light flash phenomenon observed by the crew
Definition
members after dark adaptation was attributed
to the passage of cosmic ray ions through the
The Apollo missions were the heart of NASA’s
retina of the eye (Johnston et al. 1975; Benton
manned Lunar Landing Program that took place
et al. 1977)
between 1969 and 1972 with 6 successful land-
• BIOCORE experiment during the Apollo
ings of 12 astronauts on the Moon.
17 mission that studied brain effects in pocket
mice caused by the passage of single heavy
ions (▶ HZE particles) of cosmic radiation
History (Klein 1981)
• MEED during the Apollo 16 mission that stud-
On July 20, 1969, the astronauts N. A. Armstrong ied the effects of space vacuum and solar UV
and E. E. Aldrin were the first humans to set foot radiation on different functions of microor-
on the Moon. Herewith, NASA had reached the ganisms (Taylor 1974)
ambitious goal of its manned Lunar Landing Pro-
gram. It was made possible by the strong com- The radiobiological experiments performed
mitment of the United States to manned lunar during the Apollo missions are the only ones
110 Apollo Mission
Apollo Mission, Table 1 Summary of human flights in the Apollo program to the Moon
Apollo Launch date Stay lunar
mission Mission description day/month/year surface (h) Astronauts
7 Earth orbit test 11/9/68 – Schirra,
Cunningham,
Eisele
8 Circumlunar flight 21/12/68 – Borman,
Lovell, Anders
9 Earth orbit test of LM 3/3/69 – McDivitt, Scott,
Schweickert
10 Circumlunar flight, LM separation 18/5/69 – Stafford,
Cernan, Young
11 Lunar landing, sample return 16/7/69 22.2 Armstrong,
Collins, Aldrin
12 Lunar landing, surface experiment package 14/11/69 31.5 Conrad,
Gordon, Bean
13 Lunar landing aborted 11/4/70 – Lovell, Swigert,
Haise
14 Lunar landing, highland exploration 31/1/71 33.5 Shepard,
Roosa, Mitchell
15 Lunar landing and rover, geological sampling 26/7/71 67 Scott, Worden,
Irwin
16 Lunar landing and rover, geological sampling, 16/4/72 71 Young,
Biostack and MEED experiments Mattingly,
Duke
17 Lunar landing and exploration of the Moon’s 7/12/72 75 Cernan, Evans,
geology and history, Biostack experiments Schmitt
LM lunar module
that studied the biological effects of the complete particle radiation on Apollo, Skylab, and ASTP space
interplanetary radiation field, not attenuated by missions. Nucl Track Detect 1:27–32
B€ucker H, Horneck G (1975) The biological
the Earth’s magnetic field. effectiveness of HZE-particles of cosmic
radiation studied in the Apollo
16 and 17 Biostack experiments. Acta Astronaut
See Also 2:247–264
Golombeka MP, McSween HY Jr (2007) Mars: landing
site geology, mineralogy and geochemistry. In:
▶ Biostack McFadden L-A, Weissman PR, Johnson TV (eds)
▶ Cosmic Rays in the Heliosphere Encyclopedia of the solar system, 2nd edn. Elsevier,
▶ HZE Particle Amsterdam, pp 331–348
Johnston RS, Dietlein F, Berry CA (eds) (1975) Biomedi-
▶ MEED cal results of Apollo. NASA SP-368. NASA, Wash-
▶ Microorganism ington, DC
▶ Moon, The Klein HP (1981) U.S. biological experiments in space.
▶ Radiation Biology Acta Astronaut 8:927–938
Schaber GG (2005) The U.S. geological survey,
▶ Solar UV Radiation, Biological Effects branch of astrogeology – a chronology of
▶ Space Vacuum Effects activities from conception through the end of project
Apollo (1960–1973). U.S. Department of the Interior
U.S. Geological Survey, Open-File Report
References and Further Reading 2005–1190. http://www.legislative.nasa.gov/alsj/
Schaber.html
Benton EV, Henke RP, Peterson DD (1977) Plastic Taylor G (1974) Space microbiology. Annu Rev
nuclear track detector measurements of high-LET Microbiol 28:121–137
Apsidal Angle 111
References and Further Readings

(99942) Apophis
Farnochia D, Chesley SR, Chodas PW, Micheli M, Tholen
DJ, Milani A, Elliott GT, Bernardi F (2014) A
▶ Apophis Asteroid
Yarkovsky-driven impact risk analysis for asteroid
(99942) Apophis. Icarus 224:192–200
JPL NEO Program Site Risk Page. http://neo.jpl.nasa.gov/
risk/. Last accessed 5 May 2014
NEODyS Risk Page. http://newton.dm.unipi.it/neodys/
Apophis Asteroid index.php?pc=4.0. Last accessed 5 May 2014
Sansaturio ME, Arratia O (2008) Apophis, the
Gerhard Hahn story behind the scenes. Earth Moon Planet
Asteroids and Comets, DLR, Institute of 102:425–434
Synonyms
Apparent Motion
(99942) Apophis
▶ Proper Motion
Definition
(99942) Apophis is an Aten-type asteroid, which

will make a very close approach to our planet on Apsidal Angle
April 13, 2029, passing within less than
40,000 km, close to the ring of geostationary Rory Barnes
satellites. Its size is about 375 m and its rotation Astronomy Department, University of
period 30.6 h. This close approach will change Washington, Seattle, WA, USA
the orbit substantially, from Aten type to Apollo.
History Definition
Apophis was discovered on June 19, 2004 by In planetary dynamics, the apsidal angle is the
R. A. Tucker, D. J. Tholen, and F. Bernardi at angle between the directions of closest approach
Kitt Peak. It was temporarily lost and (the apse) of two planets, as measured from
rediscovered in December 2004. Shortly thereaf- the origin of the coordinate system (usually the
ter, the close approach in 2029 was realized; even center of the star). This angle may change
a collision at that time was possible. This has with time and is coupled to the eccentricities
been ruled out based on extensive observations, of the orbits. The apsidal angle may oscillate
including radar; the orbital evolution after 2029 is about a fixed value (called apsidal libration) or
still uncertain allowing an impact probability of circulate.
5.7 106 (see impact monitoring sites at JPL
and the University of Pisa).
See Also
See Also
▶ Secular Dynamics
▶ Near-Earth Objects ▶ Secular Resonance
112 Aptamer
years later, the first RNA aptamer targeted to a

Aptamer small biomolecule was directed at ATP
(Sassanfar and Szostak 1993).
Carlos Briones Over the last two decades, a growing number
Centro de Astrobiologı́a (CSIC/INTA), of RNA and DNA aptamers have been developed
Consejo Superior de Investigaciones Cientı́ficas, against a variety of molecular targets, including
Madrid, Spain simple ions; small molecules, such as amino
acids, nucleotides, antibiotics, or metabolites;
peptides; proteins; nucleic acids; macromolecu-
Keywords lar assemblies; viruses; organelles; or even whole
cells (Klussmann 2006; Stoltenburg et al. 2007).
Molecular evolution; In vitro evolution; RNA In 2004, the first aptamer-based drug, called
world; Combinatorial nucleic acid library; Ribo- Pegaptanib, brand name Macugen, was approved
zyme; Peptide by the US Food and Drug Administration in
treatment for age-related macular degeneration
(Gragoudas et al. 2004).
Definition In parallel, peptide aptamers were obtained by
high-throughput selection methods aimed at
An aptamer (from the Latin aptus, fit, and Greek identifying members of a randomized peptide
meros, unit or part) is an in vitro selected library – usually attached at both ends to a given
oligonucleotide or peptide molecule that binds protein scaffold – by means of their interaction
to a specific target molecule. Nucleic acid with a target molecule (Colas et al. 1996). Addi-
aptamers are target-binding DNA or RNA tionally, a method termed “mRNA display”
molecules obtained by in vitro evolution. allowed the in vitro selection of peptides and
A peptide aptamer is an individual member of a proteins with the desired target-binding proper-
library of random peptide sequences that can be ties (Roberts and Szostak 1997).
selected for its ability to interact with a target
molecule.
Overview
History The preparation of a nucleic acid aptamer is cur-

rently easy and quick, since only 6–15 rounds of
By the end of the 1980s, the possibility to chem- in vitro selection or evolution are typically
ically synthesize nucleic acid pools of random required, using a combinatorial nucleic acid
sequence, as well as the availability of all the library as the starting material. Moreover, the
required enzymes for nucleic acid amplification, sensitivity and specificity of the molecular recog-
allowed the selection of target-binding RNA mol- nition between a nucleic acid aptamer and its
ecules from combinatorial nucleic acid libraries. target rival those of the antibody-antigen pairs.
The term “aptamer” was coined to denote the Additionally, different ways to increase the resis-
in vitro evolved, target-binding RNA (Ellington tance of aptamers to degradation by nuclease
and Szostak 1990), while the amplification- enzymes have been reported. These reasons,
selection process was termed “systematic evolu- together with their cost-effectiveness, make
tion of ligands by exponential enrichment” or nucleic acid aptamers not only relevant model
SELEX (Tuerk and Gold 1990). The RNA systems to address the RNA world hypothesis
aptamers selected in those two pioneering exper- (Joyce and Orgel 2006) but very useful tools in
iments were able to specifically bind different biotechnology with increasing applications in
organic dyes and a viral enzyme – the bacterio- biosensing, diagnostics, and therapy (Klussmann
phage T4 DNA polymerase – respectively. Three 2006; Mayer 2009; Germer et al. 2013).
Aptamer 113
The outcome of an in vitro evolution process is ▶ Ribozyme

usually monitored at the level of genotype, nucle- ▶ RNA
otide sequence of the evolved aptamer, and phe- ▶ RNA World A
notype, secondary/tertiary structure of the ▶ Selection
oligonucleotide, affinity and specificity of the
aptamer for its target molecule. Although nucleic
acid aptamers are artificial molecules, References and Further Reading
riboswitches have been considered “natural
aptamers” embedded in messenger RNAs since Colas P, Cohen B, Jessen T, Grishina I, McCoy J, Brent
R (1996) Genetic selection of peptide aptamers that
they act as regulatory elements for gene expres- recognize and inhibit cyclin-dependent kinase
sion by directly sensing small effector molecules 2. Nature 380:548–550
(Zhang et al. 2010). Allosteric ribozymes that Ellington AD, Szostak JW (1990) In vitro selection of
fuse one aptamer and one nucleic acid enzyme RNA molecules that bind specific ligands. Nature
346:818–822
have been developed, the ribozyme activity being Germer K, Leonard M, Zhang X (2013) RNA aptamers
modulated by the binding of an effector molecule and their therapeutic and diagnostic applications. Int
to the aptamer domain. The biotechnological J Biochem Mol Biol 4:27–40
applications of such “aptazymes” – also known Gragoudas ES, Adamis AP, Cunningham ET Jr,
Feinsod M, Guyer DR, VEGF Inhibition Study in
as artificial riboswitches – are increasingly rec- Ocular Neovascularization Clinical Trial
ognized (Wieland and Hartig 2008). Recently, Group (2004) Pegaptanib for neovascular
the unveiled ability of certain nucleic acid ana- age-related macular degeneration. N Engl J Med
logues to fold into three-dimensional structures 351:2805–2816
Joyce GF, Orgel LE (2006) Progress toward understand-
allowed the in vitro evolution of aptamers har- ing the origin of the RNA world. In: Gesteland RF,
boring different molecular scaffolds, some of Cech TR, Atkins JF (eds) The RNA world. Cold
which bind their targets with an affinity similar Spring Harbor Laboratory Press, New York
to that of RNA or DNA aptamers (Pinheiro Klussmann S (ed) (2006) The aptamer handbook. Wiley-
VCH, Weinheim
et al. 2012). Li J, Tan S, Chen X, Zhang CY, Zhang Y (2011) Peptide
In addition to nucleic acid aptamers, the pep- aptamers with biological and therapeutic applications.
tide aptamer approach has utilized small inert Curr Med Chem 18:4215–4222
scaffold proteins to expose the randomizable pep- Mayer G (ed) (2009) Nucleic acid and peptide aptamers.
Methods and protocols. Springer, Heidelberg
tide region. Alternatively, the mRNA display Pinheiro VB, Taylor AI, Cozens C, Abramov M,
technique has been used to develop peptide Renders M, Zhang S, Chaput JC, Wengel J, Peak-
aptamers, and “stand-alone” interfering peptides Chew SY, McLaughlin SH, Herdewijn P, Holliger
have also been selected. The ability of peptide P (2012) Synthetic genetic polymers capable of hered-
ity and evolution. Science 336:341–344
aptamers to specifically interact with different Roberts RW, Szostak JW (1997) RNA-peptide fusions for
proteins and other target molecules offers broad the in vitro selection of peptides and proteins. Proc
applicability in biochemistry and biomedicine Natl Acad Sci U S A 94:12297–12302
(Mayer 2009; Li et al. 2011). Sassanfar M, Szostak JW (1993) An RNA motif that binds
ATP. Nature 364:550–553
Stoltenburg R, Reinemann C, Strehlitz B (2007)
SELEX – a (r)evolutionary method to generate high-
See Also affinity nucleic acid ligands. Biomol Eng 24:381–403
Tuerk C, Gold L (1990) Systematic evolution of
ligands by exponential enrichment: RNA ligands to
▶ Amplification (Genetics) bacteriophage T4 DNA polymerase. Science
▶ Aptasensor 249:505–510
▶ Combinatorial Nucleic Acid Library Wieland M, Hartig JS (2008) Artificial riboswitches: syn-
▶ DNA thetic mRNA-based regulators of gene expression.
Chembiochem 9:1873–1878
▶ Evolution, In Vitro Zhang J, Lau MW, Ferré-D’Amaré AR (2010) Ribozymes
▶ Evolution, Molecular and riboswitches: modulation of RNA function by
▶ Peptide small molecules. Biochemistry 49:9123–9131
114 Aptamer-Based Biosensor
acid molecules (Tuerk and Gold 1990; Ellington

Aptamer-Based Biosensor and Szostak 1990). Adapting both technologies,
aptamers-based biosensors were established in
▶ Aptasensor 1996 when Drolet and coworkers (Drolet
et al. 1996) described a modification of the
renowned enzyme-linked immunosorbent assay
(ELISA) which utilized an aptamer instead an
Aptasensor antibody as biorecognition element. The assay
was named “enzyme-linked oligonucleotide
Miguel Moreno assay” (ELONA), and this evidence opened the
Centro de Astrobiologı́a, CSIC, Madrid, Spain door to novel ways of biosensing using nucleic
acids to detect a wide range of molecules in a
variety of formats.
Keywords
Aptamer; Biosensor; DNA; RNA Overview
Aptamers are attracting interest in the areas of

Synonyms therapeutics and diagnostics and offer themselves
as ideal candidates for use as the recognition
Aptamer-based biosensor elements in biosensors since they possess many
advantages over the state-of-the-art affinity sen-
sors. Aptasensors show very high sensitivity,
Definition specificity, and reproducibility against a wide
variety of targets. Thus, they are rapidly emerg-
An aptasensor is a particular class of biosensor ing as promising candidates for high-throughput
where the biological recognition element is a analytical methods that have to deal with tiny
DNA or RNA aptamer. In an aptasensor, the quantities of the queried analytes. Analogous to
aptamer recognizes the molecular target against immunoassays (those based on the antigen-
which it was previously in vitro selected. The antibody interaction), aptamer-based bioassays
aptamer-target reaction is independent of both can adopt different configurations to transduce
the type of detection system and the kind of biorecognition events, including aptamer-based
transducer employed. Aptasensors can be easily microarrays (Collett et al. 2005), aptamer-capped
multiplexed to detect a variety of aptamer-target gold nanoparticles (Song et al. 2012), quantum-
reactions simultaneously. dot aptamer conjugates (Levy et al. 2005), and
electrical (Willner and Zayats 2007) and electro-
chemical (Moreno et al. 2011) aptasensors,
History among many others (Citartan et al. 2012).
Aptamers offer an extensive range of advan-
Since the development of the first glucose bio- tages over other existing biological recognition
sensor in 1962 (Clark and Lions 1962), an exten- elements in terms of stability, design flexibility,
sive choice of biosensors has been developed. and cost-effectiveness. To name a few, aptamers
Among them, nucleic acid-based biosensors are can bind to their targets with affinities and spec-
of particular interest due to their practical appli- ificities equivalent to those of monoclonal anti-
cations in different fields of genomic research. In bodies and can be selected to bind a wide range of
1990 two independent groups simultaneously targets including those that are toxic or not inher-
described aptamers as target-binding nucleic ently immunogenic. Additionally, the affinities
Aqueous Interfaces 115
and specificities of aptamers can be easily tai- References and Further Reading
lored (in contrast to those of antibodies) and can
be more readily engineered than antibodies for Cho EJ, Lee J-W, Ellington AD (2009) Applications
of aptamers as sensors. Annu Rev Anal Chem A
their use as biosensing elements. Finally,
2:241–264
aptamers can be synthesized, chemically modi- Citartan M, Gopinath SCB, Tominaga J, Tan S-C, Tang
fied, and stored until needed and are resistant to T-H (2012) Assays for aptamer-based platforms.
denaturation and degradation. The in-depth Biosens Bioelectron 34:1–11
Clark LC, Lions C (1962) Electrode systems for continu-
knowledge of aptamer conformational properties
ous monitoring in cardiovascular surgery. Ann Acad
and ligand-binding mechanisms has triggered Sci 102:29
profound attention among researchers for devel- Collett JR, Cho EJ, Ellington AD (2005) Production and
oping aptasensor bioassays, as reflected in the processing of aptamer microarrays. Methods 37:4–15
Drolet DW, Moon-McDermott L, Romig TS (1996) An
exponential increase of published articles
enzyme-linked oligonucleotide assay. Nat Biotechnol
(Citartan et al. 2012). 14:1021–1025
Nevertheless, aptasensors compete with other Ellington AD, Szostak JW (1990) In vitro selection of
well-established biosensors, essentially RNA molecules that bind specific ligands. Nature
346:818–822
antibodies-based ones, and they have been
Fischer NO, Tarasow TM, Tok JBH (2007) Aptasensors
mainly focused on the applications where anti- for biosecurity applications. Curr Opin Chem Biol
bodies cannot achieve the desired goals. The 11:316–328
most straightforward application of aptasensors Levy M, Cater SF, Ellington AD (2005) Quantum-dot
aptamer beacons for the detection of proteins.
lies in the fields of food safety testing and envi-
Chembiochem 6:2163–2166
ronmental pollution control. Thus, aptasensors Moreno M, Gonzalez VM, Rincon E, Domingo A,
developed for the detection of small molecule Dominguez E (2011) Aptasensor based on the selec-
contaminants including antibiotics, toxins, pesti- tive electrodeposition of protein-linked gold
nanoparticles on screen-printed electrodes. Analyst
cides, and heavy metals (that may be present in a
136:1810–1815
wide variety of food products and environmental Song K-M, Jeong E, Jeon W, Cho M, Ban C (2012)
samples) are of particular interest (Cho Aptasensor for ampicillin using gold nanoparticle
et al. 2009; Fischer et al. 2007). based dual fluorescence–colorimetric methods. Anal
Bioanal Chem 402:2153–2161
In astrobiology, aptasensors are called to be a
functional tool for the detection of both mole- ligands by exponential enrichment: RNA ligands to
cules with limited antigenicity present in differ- bacteriophage T4 DNA polymerase. Science
ent extreme environments and biosignatures of 249:505–510
Willner I, Zayats M (2007) Electronic aptamer-based sen-
extinct or extant life in planetary exploration. In
sors. Angew Chem Int Ed 46:6408–6418
the field of biomarker detection in space mis-
sions, robust and very stable aptasensors might
be developed with extended capabilities to over-
come the extreme conditions of long travel time
and planetary exploration.
Aqueous Interfaces
See Also Veronica Vaida and Elizabeth C. Griffith

University of Colorado, Boulder, CO, USA
▶ Antibody
▶ Aptamer
▶ Biomarkers Keywords
▶ Biosensor
▶ Combinatorial Nucleic Acid Library Water surface; Unique reaction environment;
▶ Evolution, In Vitro Amphiphilic molecules; Surfactant film
116 Aqueous Interfaces
Definition Overview
An aqueous interface is the dividing surface Aqueous interfaces are found throughout nature:
between two media, one of which is water. at the surface of lakes and oceans, at the surface
of atmospheric aerosol particles, or even at the
interface between a membrane and its surround-
History ing bulk water environment. Air-aqueous inter-
faces are particularly interesting in an
Atmospheric interfaces including rocks and clays astrobiological context due to their provision of
have long been considered in the origin of life a unique reaction environment, having the ability
scenarios. Aqueous interfaces (water surfaces) as to concentrate and align surface active molecules
found on oceans, lakes, rivers, and atmospheric (Goldacre 1958; Shah 1972). It was pointed
aerosols were first suggested by Goldacre (1958) out that the vast collective surface area of
to be interesting in a prebiotic context due to their atmospheric aerosols provides diverse and fluc-
ability to concentrate organic molecules and sub- tuating environments for chemistry and is appli-
sequently fold and pinch off into enclosures rem- cable to any rotating planetary body with a liquid
iniscent of cells. The use of Global atmospheric ocean (Dobson et al. 2000; Griffith et al. 2012).
aerosols were pointed out in this context (Shah A planet rotating on a tilted axis results in thermal
1972) and suggested as effective prebiotic and pressure gradients that produce wind.
microreactors in different contexts later This wind acting on a liquid ocean produces
(Lerman 2010; Dobson et al. 2000; Tverdislov sea spray from which aqueous atmospheric aero-
and Yakovenko 2008). In addition, aqueous inter- sols are born (as depicted in the Fig. 1). Any
faces have recently been suggested as favorable organic material residing at or near the ocean
environments for biomolecular synthesis that is surface will be entrained in these aerosols and
difficult or impossible in the bulk ocean (Dobson can partition to their surface (an aqueous inter-
et al. 2000; Ruiz-Bermejo et al. 2010; Griffith face), forming a surfactant film around an aque-
et al. 2012). ous core.
Aqueous Interfaces,
Fig. 1 Depiction of birth
of aerosols from sea spray
as well as schematic
representation of peptide
bond formation at the
air-aqueous interface from
amino acid (AA) precursors
Aquifer (Mars) 117
These interfaces allow for concentration of Lynden-Bell RM, Morris SC, Barrow JD, Finney JL,
reactant species over the bulk aqueous solution, Harper CL Jr (eds) Water and life: the unique proper-
ties of water. CRC Press, Boca Raton, pp 259–290
alteration of the ionization state of surface- Lynden-Bell RM, Morris SC, Barrow JD, Finney JL, A
residing reactants, and orientation of amphiphilic Harper CL Jr (eds) (2010) Water and life: the unique
molecules, and they can even promote chemistry properties of water. CRC Press, Boca Raton
that is inaccessible in bulk water (Griffith and Pratt LR, Pohorille A (2002) Hydrophobic effects and
modeling of biophysical aqueous solution interfaces.
Vaida 2013). One example of a key reaction Chem Rev 102(8):2671–2691
made possible by an aqueous interface is the Ruiz-Bermejo M, Menor-Salvan C, Zorzano MP,
formation of peptide bonds in the absence of El-Hachemi Z, Osuna-Esteban S, Veintemillas-
enzymatic catalysis (Griffith and Vaida 2012) Verdaguer S (2010) Water interfacial processes in
prebiotic chemistry. In: Hegedus S, Csonka J (eds)
illustrated schematically in Fig. 1. Peptide Astrobiology: physical origin, biological evolution
bonds are a key bond in modern biology as they and spatial distribution. Nova Science Publishers
are the link between amino acid building blocks Inc., New York, pp 27–57
in proteins (one of the three principle biopoly- Shah DO (1972) The origin of membranes and related
surface phenomena. Exobiology. C. Ponnamperuma.
mers along with RNA and DNA). In addition, North-Holland Publishing Co., Amsterdam,
theoretical and experimental studies have been pp 235–265
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of bimolecular assemblies at the surface of liquid origin of life. Surv Geophys 23(5):379–409
Tverdislov VA, Yakovenko LV (2008) Physical aspects of
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(Pratt and Pohorille 2002). chiral asymmetries as two fundamental asymmetry
types. Mosc Univ Phys Bull 63(3):151–163
Cross-References
▶ Aerosols Aquifer (Mars)

▶ Amino Acid
▶ Amphiphile Alessandro Airo
▶ Membrane Institut f€ur Geologische Wissenschaften
▶ Water, Solvent of Life Tektonik und Sedimentäre Geologie, Freie
Universität Berlin, Fachbereich
Dobson CM, Ellison GB, Tuck AF, Vaida V (2000) Atmo-

spheric aerosols as prebiotic chemical reactors. Proc
Definition
Natl Acad Sci U S A 97(22):11864–11868
Goldacre RJ (1958) Surface films, their collapse on com- It is widely accepted that liquid water was present
pression, the shape and size of cells and the origin of on the surface of Mars during its early history.
life. In: Danielli JF, Parkhurst KGA, Riddiford AC
(eds) Surface phenomena in chemistry and biology.
Although parts of this water have vanished into
Pergamon Press, New York, pp 12–27 space or were consumed in chemical reactions,
Griffith EC, Vaida V (2012) In situ observation of peptide substantial amounts are still present today. The
bond formation at the water-air interface. Proc Natl current physical conditions on Mars’ surface usu-
Acad Sci U S A 109(39):15697–15701
Griffith EC, Vaida V (2013) Ionization state of
ally do not allow liquid water to be stable, and
L-phenylalanine at the air-water interface. J Am therefore, it occurs as water ice within the pore
Chem Soc 135(2):710–716 space of the permafrost soil or as few km-thick
Griffith EC, Tuck AF, Vaida V (2012) Ocean-atmosphere polar ice caps. It can be assumed that at a certain
interactions in the emergence of complexity in simple
chemical systems. Acc Chem Res 45(12):2106–2113
depth below the surface, usually estimated to be a
Lerman L (2010) The primordial bubble: water, few kilometers, the cryosphere transitions into an
symmetry-breaking, and the origin of life. In: aquifer (Lasue et al. 2013). The depth at which
118 Arachnoid
the water ice turns into groundwater depends on See Also

not well known factors, such as the water salinity,
the soil porosity and permeability, or the geother- ▶ Venus
mal gradient and thermal conductivity of the sub-
surface. The presence of groundwater on Mars is
insofar of interest to astrobiologists as aquifers on
Earth are populated by microbial life down to a
Archaea
depth of a few kilometers (Michalski et al. 2013).
Antonio Ventosa and Rafael R. de la Haba
Department of Microbiology and Parasitology,
See Also
Faculty of Pharmacy, University of Sevilla,
Sevilla, Spain
▶ Crater Lakes (Mars)
▶ Gullies
▶ Habitability on Mars
Keywords
▶ Heat Transfer, Planetary
▶ Outflow Channels
Domain; Evolution; Extremophiles; Molecular
▶ Polar Caps (Mars)
adaptation; Phylogeny; 16S rRNA sequencing

Synonyms
Lasue J, Mangold N, Hauber E, Clifford S, Feldman W,
Gasnault O, Grima C, Maurice S, Mousis O (2013)
Archaeobacteria
Quantitative assessment of the Martian hydrosphere.
Space Sci Rev 174:155–212
Michalski JR, Cuadros J, Niles PB, Parnell J, Rogers AD,
Wright SP (2013) Groundwater activity on Mars and Definition
implications for a deep biosphere. Nat Geosci
6:133–138
The Archaea are a phylogenetically coherent
group of prokaryotes that have a different orga-
nization than the ▶ Bacteria.
Arachnoid
Jörn Helbert History

DLR, Institut f€
ur Planetenforschung, Berlin,
Germany Woese and Fox (1977) proposed that prokaryotes
were not a monophyletic group. Based on the
comparison of their small subunit ribosomal
Definition RNA sequences, the prokaryotes comprise two
distinct evolutionary lineages that are represented
An Arachnoid is a type of landform only seen on by the ▶ Bacteria and the Archaea (that formerly
the surface of ▶ Venus, which is believed to have were designated as Archaebacteria by Woese
a volcanic origin. Arachnoids get their name from et al. 1978). The concept of a third domain of
their resemblance to spider webs. They appear as life, which explained several structural, meta-
concentric ovals surrounded by a complex net- bolic, and molecular differences with respect to
work of fractures, and are spanned up to 200 km. other prokaryotes, was initially poorly accepted
Over 30 arachnoids have been identified on by the scientific community. However, the con-
Venus, so far. cept of the Archaea was advanced through
Archaea 119
studies and meetings carried out by O. Kandler, units; (4) although the translation machinery of
W. Zillig, and K.O. Stetter, among others. The Archaea is similar to that of bacteria (70S ribo-
Archaea have similarities and are considered somes with 50S and 30S subunits, similar length A
phylogenetically more closely related with the ribosomal RNAs, transcriptional and transla-
▶ Eukarya (Woese et al. 1990). tional coupling, etc.), there are an important num-
ber of specific features not present in Bacteria,
some of which are specific to Archaea while
Overview others are similar to Eukarya. For example,
almost all antibiotics that inhibit bacterial trans-
Archaea have distinct molecular characteristics lation are ineffective in the Archaea; Bacteria use
that clearly distinguish them from the Bacteria N-formyl-methionyl-tRNA for translational start
and the Eukarya, and evolutionary studies have codons, while Archaea use unmodified initiator
highlighted their role on the development of life methionine in translation, similar to Eukarya.
on our planet. Archaea have been associated with Besides, Archaea and Eukarya share a common
▶ extreme environments and many of them are characteristic, elongation factor 2 (EF-2), which
extremophilic microorganisms, showing interest- is ADP-ribosylated by diphtherial toxin.
ing characteristics and applications for industrial The Archaea are subdivided into five phyla, of
and other purposes. Many of them are considered which two, the Crenarchaeota and the
to be microorganisms that are able to grow on the Euryarchaeota, are most extensively studied.
limits of life. Their ability to thrive in extreme The classification of Archaea has been widely
environments has expanded the horizons for discussed and several proposals have been published.
Astrobiology as they are considered counterparts The most widely accepted include the Archaea
for extraterrestrial life. as a higher taxon with the range of “Domain,”
The Archaea are characterized by a cellular which includes the following five phyla:
morphology similar to those of most Bacteria Crenarchaeota, Euryarchaeota, Korarchaeota,
(rods, cocci, irregular cells, etc.). However, Nanoarchaeota, and Thaumarchaeota (Parte 2015).
myceliar or multicellular stages with cellular dif- The phylum Crenarchaeota includes a single class,
ferentiation have not been described. On the con- Thermoprotei, with five orders: Acidilobales,
trary, unique morphologies have been described Desulfurococcales, Fervidicoccales, Sulfolobales,
for some Archaea, such as the square flat cells of and Thermoproteales. The phylum Euryarchaeota
some haloarchaea (Haloquadratum walsbyi) or includes eight classes: Archaeoglobi, Halobacteria,
amoeba-like cells (Thermoplasma and other Methanobacteria, Methanococci, Methanomicrobia,
microorganisms). Other characteristics of the Methanopyri, Thermococci, and Thermoplasmata.
Archaea that define their differential status with The phylum Korarchaeota includes nonculti-
respect to the other living organisms are: (1) the vated Archaea designated as “Candidatus
presence of phytanyl ether instead of fatty acid Korarchaeum,” the phylum Nanoarchaeota
ester lipids in their membranes; (2) the absence of includes the genus “Nanoarchaeum,” and finally,
peptidoglycan (murein) in their cell walls and a the phylum Thaumarchaeota includes the genus
frequent presence of proteinaceous S-layers (only “Cenarchaeum.”
a few have a polysaccharide cell wall), as well as The phylum Euryarchaeota includes two of
the absence of a periplasmic space; (3) their com- the most typical groups that were identified in
plex DNA-dependent RNA polymerases (early the early studies by Woese and coworkers as
in vitro studies using several inhibitors showed members of the Archaea: the ▶ methanogens
that the transcription machinery in Archaea is and the haloarchaea (also designated as
more closely related to that of Eukarya than to halobacteria). The methanogenic Archaea are
Bacteria); the sequences of the archaeal RNA anaerobic organisms that produce methane as
polymerases resemble some eukaryotic RNA the major end product of their metabolism. Phy-
polymerases and consist of up to 13 different logenetically methanogens are very diverse and
120 Archaea
are represented by a large number of species alkaline pH and inhabiting soda lakes. Other typ-
belonging to many genera, grouped in 12 families ical features of haloarchaea are: their production
within 6 orders. They are found on a variety of of red- to pink-pigmented colonies due to the
anoxic environments such as ocean and lake sed- presence of bacterioruberins (C50 carotenoids),
iments, hydrothermal vents, animal digestive although there are a few exceptions; the presence,
tracts, anaerobic sludge digesters, etc. The typical in some of them, of retinal-based pigments
growth compounds of methanogens are H2 and (bacteriorhodopsin), that act as a proton pump
CO2, or short-chain (C1-C5) organic compounds driven by light energy; or the presence of typical
(formate, acetate, ethanol, trimethylamine, etc.). archaeal polar lipids, with ether-linked
H2 is used as electron donor for CO2 reduction, phosphoglycerides that can be easily detected
and electrons can also be derived from by thin-layer chromatography (a feature that is
formate, CO, or specific alcohols. Among the widely used for the taxonomic differentiation of
microorganisms that have been used as models most genera of haloarchaea) (Grant et al. 2001).
for studying methanogenesis are species of the Haloarchaea are excellent models for the
genera Methanobacterium, Methanother- study of the molecular biology and other struc-
mobacter, Methanobrevibacter, Methanosarcina, tural features of Archaea, as well as their mecha-
Methanococcus, among others (Dworkin nisms of adaptation to extreme conditions of
et al. 2002; Madigan et al. 2008). salinity, alkaline pH, and moderate temperature,
Haloarchaea are represented by a group of and several species have been used for such
extremely halophilic aerobic Archaea, which tax- purposes due to their ease of manipulation
onomically are placed within a single class, under laboratory conditions: they grow in complex
Halobacteria, order Halobacteriales, family media (with the appropriate salt content) under aer-
Halobacteriaceae (Grant et al. 2001). Currently obic conditions using the standard procedures uti-
they are represented by almost 50 genera and a lized for most nonfastidious prokaryotes. Some
large number of species that are characterized by species used for such studies include Halobacterium
their Na+ requirements. They are considered to be salinarum, Haloarcula marismortui, Haloferax
organisms that are able to grow under higher salt volcanii, and more recently, the square
concentrations, in saturated NaCl habitats. Their haloarchaeon Haloquadratum walsbyi (recently iso-
optimal NaCl requirements are in the range lated and referred to as “Walsby’s square bacte-
3.5–4.5 M NaCl and they are not able to grow in rium”). In addition, several biotechnological
media without NaCl, thus, they have a specific applications have been suggested, such as the com-
requirement for NaCl, which has led to detailed mercial production of bacteriorhodopsin, the pro-
studies of their mechanisms of haloadaptation. In duction of extracellular hydrolytic enzymes or
contrast to most other prokaryotes, which accu- exopolysaccharides, the use of polyhydrox-
mulate intracellular organic compounds desig- yalkanoates (PHAs) as bioplastics, or the production
nated as ▶ compatible solutes, haloarchaea of halocins (archaeocins, proteinaceous archaeal
compensate for the high salt concentration in antimicrobials).
the environment by accumulating ions, mainly With a few bacterial exceptions, most
up to 5 M KCl. They are normal inhabitants of ▶ hyperthermophiles (defined as organisms
hypersaline environments, being the predominant showing optimal growth at 80 C or higher) are
microbiota of saturated ponds of salterns and salt species of Archaea. They are inhabitants of
lakes (they may reach high cell densities, hot springs, solfataric and volcanic areas,
>107 cell ml1); they are also found in salt or deep-subsurface aquifers, submarine vents
salted products (salted fish or meats, salted (“black smokers”), etc. Hyperthermophiles
fermented foods), salt deposits (mines), salted include several methanogens, as well as
hides, and saline soils. Most haloarchaea grow members of a variety of genera of the
at neutral pH values but some species are Archaeoglobales, Thermococcales, Desulfuro-
haloalkaliphilic, being able to grow optimally at coccales, Thermoproteales, or Sulfolabales.
Archaea 121
They are excellent models for the study of the References and Further Reading
metabolisms of sulfur and inorganic sulfur com-
pounds; many species use inorganic sulfur com- Blum P (ed) (2001) Archaea: ancient microbes, extreme
environments, and the origin of life. Academic, San A
pounds as electron acceptors or donors. Some of
Diego
the most hyperthermophilic organisms known are Blum P (ed) (2008) Archaea: new models for prokaryotic
Pyrolobus fumarii (optimal growth at 106 C, biology. Caister Academic Press, Norfolk
range 90–113 C), Pyrodictium occultum Boone DR, Castenholz RW, Garrity GM (2001) Bergey’s
manual of systematic bacteriology, 2nd edn. The
(optimal growth at 105 C, range 85–110 C),
Archaea and the deeply branching and phototrophic
Pyrococcus furiosus and Pyrococcus woesei Bacteria, vol 1. Springer, New York
(optimal growth at 100–103 C, range Brochier-Armanet C, Boussau B, Gribaldo S, Forterre
70–105 C), Pyrobaculum aerophylum (optimal P (2008) Mesophilic crenarchaeota: proposal for a
third archaeal phylum, the Thaumarchaeota. Nat Rev
growth at 100 C, range 75–104 C), and Microbiol 6:245–252
Pyrobaculum islandicum (optimal growth at 100 Casanueva A, Galada N, Baker GC, Grant WD, Heaphy S,

C, range 74–102 C). Jones B, Yanhe M, Ventosa A, Blamey J, Cowan DA
The phylum Nanoarchaeota is known for a (2008) Nanoarchaeal 16S rRNA gene sequences are
widely dispersed in hyperthermophilic and mesophilic
single species, Nanoarchaeum equitans, to date
halophilic environments. Extremophiles 12:651–656
a hyperthermophilic archaeon that lives in a sym- Cavicchioli R (ed) (2007) Archaea: molecular and cellular
biotic association with the Crenarchaeote biology. ASM Press, Washington, DC
Ignicoccus, a sulfur-dependent anaerobic hyper- Dworkin M, Falkow S, Rosenberg E, Schleifer K-H,
Stackebrandt E (eds) (2002) The prokaryotes: an
thermophile. The cells are spherical and only evolving electronic resource for the microbiological
about 400 nm in diameter; they grow attached to community, 3rd edn, release 3.19 ed. Springer, New
the surface of a specific archaeal host (Hubber York. http://link.springer-ny.com/link/service/books/
et al. 2002). This archaeon was isolated from a 10125/
Garret RA, Klenk H-P (eds) (2007) Archaea:
submarine hot vent, but recent studies have evolution, physiology and molecular biology. Black-
shown that nanoarchaea may be widely dispersed well, Oxford
in hyperthermophilic and mesophilic halophilic Grant WD, Kamekura M, McGenity TJ, Ventosa A (2001)
environments (Casanueva et al. 2008; Rinke et al. Class III. Halobacteria class. In: Boone DR,
Castenholz RW, Garrity GM (eds) Bergey’s manual
2013).
of systematic bacteriology, 2nd edn. The Archaea and
the deeply branching and phototrophic Bacteria,
vol 1. Springer, New York
Hubber H, Hohn MJ, Rachel R, Fuchs T, Wimmer VC,
Stetter KO (2002) A new phylum of archaea
See Also represented by a nanosized hyperthermophilic symbi-
ont. Nature 417:63–67
▶ Bacteria Kates M, Kushner DJ, Matheson AT (1993) The biochem-
▶ Compatible Solute istry of Archaea (Archaebacteria). Elsevier,
Amsterdam
▶ Crenarchaeota
Madigan MT, Martinko JM, Dunlap PV, Clark DP
▶ Domain (Taxonomy) (2008) Brock biology of microorganisms, 12th edn.
▶ Eukarya Benjamin Cummings, San Francisco
▶ Euryarchaeota Parte A (2015) List of prokaryotic names with standing in
nomenclature. http://www.bacterio.cict.fr/.
Pfeifer F, Palm P, Schleifer K-H (1994) Molecular biology
▶ Halophile of Archaea. Gustav Fischer Verlag, Stuttgart
▶ Hyperthermophile Rinke C, Schwientek P, Sczyrba A, Ivanova NN, Ander-
▶ Korarchaeota son IJ, Cheng JF, Darling A, Malfatti S, Swan BK,
Gies EA, Dodsworth JA, Hedlund BP, Tsiamis G,
▶ Membrane
Sievert SM, Liu WT, Eisen JA, Hallam SJ, Kyrpides
▶ Methanogens NC, Stepanauskas R, Rubin EM, Hugenholtz P,
▶ Nanoarchaeota Woyke T (2013) Insights into the phylogeny and cod-
▶ Phylogenetic Tree ing potential of microbial dark matter. Nature
499:431–437
▶ Prokaryote
122 Archaeobacteria
Robb FT, Place AR, Sowers KR, Schreier HJ, surface. Astrobiology-related studies such as
DasSarma S, Fleischmann EM (eds) (1995) Archaea: search of pristine morphological or chemical
a laboratory manual. Cold Spring Harbor, New York
Ventosa A (2006) Unusual micro-organisms from unusual traces of early life form an important part of
habitats: hypersaline environments. In: Logan NA, these projects.
Lappin-Scott HM, Oyston PCF (eds) Prokaryotic
diversity: mechanisms and significance. Cambridge
University Press, Cambridge
Woese CR, Fox GE (1977) The phylogenetic structure of Overview
the prokaryotic domain: the primary kingdoms. Proc
Natl Acad Sci U S A 74:5088–5090 To date, both volcanic and sedimentary
Woese CR, Wolfe RS (eds) (1985) The bacteria: a treatise sequences have been targeted, and the recovered
on structure and function. Archaeabacteria, vol VII-
I. Academic, New York cores have been analyzed to investigate condi-
Woese CR, Magrum LJ, Fox GE (1978) Archaebacteria. tions at the surface of the Archean Earth – the
J Mol Evol 11:245–251 composition, temperature, and redox state of the
Woese CR, Kandler O, Wheelis ML (1990) Towards a Archean ocean and atmosphere and the volcanic
natural system of organisms: proposal for the domains
Archaea, Bacteria and Eukarya. Proc Natl Acad Sci and sedimentary processes that operated early in
U S A 87:4576–4579 Earth history – and, above all, to search for evi-
dence of primitive life. The focus has been the
▶ Pilbara Craton in Western Australia, where
four separate programs have been carried out,
Archaeobacteria each involving collaboration between geologists
from Australian universities, the Western Austra-
▶ Archaea lian Geological Survey, and foreign agencies.
The four programs are (1) the Archean Biosphere
Drilling Project (ABDP) cosponsored by several
Japanese Universities, (2) the Deep Time Drilling
Project (DTDP) of the NASA Astrobiology Insti-
Archean Biosignatures tute, (3) the Pilbara Drilling Project (PDP) of IPG
Paris, and (4) Dixon Island-Cleaverville Drilling
▶ Archean Traces of Life Project (DXCL-DP) supported by the Japanese
Ministry of Education, Culture, Sports, Science,
and Technology (MEXT).
In South Africa’s ▶ Barberton Greenstone
Archean Drilling Projects Belt, two drilling projects were completed to date:
Nicholas Arndt The Barberton Barite Drilling Project (CNRS,

ISTerre, Université Grenoble Alpes, France IPGP) had the objective to obtain a representa-
tive sequence of black cherts, shales, tuffaceous
sandstones and siltstones, jasper deposits, and
Definition bedded barite which is a conspicuous assem-
blage of rock types typically observed from
There are several scientific drilling projects that Early Archean seafloor hydrothermal settings.
have been carried out in Archean terrains in the They well penetrated a section in the west limb
past decade and others that are planned in the near of the Baryte Syncline to a depth of 182 m.
future. The aim of most of these drilling projects The Barberton Drilling Project (ICDP; July
is to recover relatively well-preserved rock sam- 2011–May 2012) included five diamond-core
ples from below the present weathering profile holes, each several hundred meters long.
and to obtain continuous rock cores that retain BARB-1 and BARB-2 targeted komatiitic
soft or friable units that outcrop poorly at the rocks near Tjakastad in the southern part of the
Archean Environmental Conditions 123
belt, whereas BARB-3 drilled 899.5 m through conditions of the surface of the Earth during the
the Buck Reef Chert, both of the ▶ Onverwacht ▶ Archean eon. The surface of the Archean Earth
Group. BARB-4 targeted turbidites and banded- was in many ways similar to that of today. Oceans A
iron formation and BARB-5 ▶ barite, turbidites, likely covered most of the globe, but there were
and ▶ spherules, both of the Mapepe Formation also regions of dry land. However, the oceanic
(lower Fig Tree Group). A total of 3,052 m of crust was almost as thick as the ▶ continental
core was recovered. crust, mountain ranges were not very high, and
parts of oceanic ridges and plateaus (thick piles of
Two programs focused on the Archean- flat-lying lava flows) were emergent. Geological
Proterozoic transition. A series of short holes processes such as volcanism, erosion, and sedi-
have been drilled in Russian Fennoscandia to ment deposition operated but were influenced by
sample the 500-million-year interval defining a lack of vegetation, higher ocean temperatures,
the Archean-Paleoproterozoic transition; the different water composition, and a hotter, more
Agouron Griqualand Paleoproterozoic Drilling aggressive, acidic atmosphere. In addition,
Project straddled a similar interval in the North- coastal settings may have been subjected to
ern Cape province of South Africa. more intense reworking by high and more fre-
quent tides.
See Also
Overview
▶ Archean Traces of Life
The surface of the Archean Earth was in many
▶ Barberton Greenstone Belt
ways similar to that of today. Oceans covered
▶ Early Archean
most of the globe, but there were also regions of
▶ Microfossils
dry land. The total area covered by oceans was
▶ Pilbara Craton
greater than now for three reasons. First, the
▶ Proterozoic Eon
volume of continental crust may have been less,
if continental crust indeed grew (progressively or
in spurts) through time (Benn et al. 2006).
Second, the oceans might have been more
Archean Environmental Conditions voluminous because high temperatures in the
mantle (Nisbet et al. 1993) destabilized
Christoph Heubeck1 and Nicholas Arndt2 hydrous minerals and drove water to the
1
Institut f€
ur Geowissenschaften, Friedrich- surface. Third, oceanic crust was thicker. Its
Schiller-Universität Jena, Jena, Germany top lay at a shallower depth than today’s
2
ISTerre, Université Grenoble Alpes, France sea floor, thus displacing water which
inundated low-lying margins of continents
(Eriksson 1999).
Keywords Mountain ranges existed but were not as high
as those of today because the continental crust
Chert; Komatiite; Oceans; Sediment; Traces of was heated internally and rendered more ductile
life; Carbonates by more abundant radioactive elements. Conti-
nental crust was relatively thin, while oceanic
crust, produced by high-degree melting of the
Definition hotter mantle, was far thicker (Sleep and Windley
1982). The subdued topography, the limited con-
The term “Archean environmental conditions” trast between the thicknesses of oceanic and con-
refers to the geological, physical, and chemical tinental crust, combined with bigger oceans,
124 Archean Environmental Conditions
meant that much of the continental crust was barren shorelines in much wider tidal belts than
flooded (Arndt 1998). we see today.
Just as during more recent geological The oceanic crust was composed of basaltic
history, global temperatures waxed and waned. lavas like that of modern crust, but was more
The world’s oldest glaciation is reported magnesian (picritic) in places (Sleep and
from the 2.9 Ga-old Mozaan Group on South Windley 1982). Parts of mid-ocean ridges and
Africa’s Kaapvaal Craton (Young et al. 1998) the summits of oceanic plateaus may have been
but on the whole temperatures appear to have emergent, forming what might be called
been clement or high. The O and Si isotopic “melano- (dark-colored) continents.” The pelagic
compositions of Archean ▶ cherts suggest sediment that covered this crust was different
that ocean temperatures were commonly above from that of today. An absence of shell-forming
40 C and possibly as high as 80 C (Knauth and organisms precluded the formation of biogenic
Lowe 2003). The atmosphere contained very calcareous or siliceous oozes; in their place
little or no free oxygen but was richer in CO2; occurred Si- or Fe-rich sediments that precipi-
rainwater was thus somewhat acid. Due to the tated directly from the high-temperature seawater
absence of a protective ozone shield, UV flux at that contained high concentrations of these ele-
the Earth’s surface was high. The atmosphere ments. Depending on the intensity of weathering
also contained SO and SO2, causing sulfuric of mafic material on exposed regions, much clay
acid haze; haze from nitrous oxides and may have been produced, washed into the sea and
organics likely occurred as well. The normal deposited on oceanic crust. Hydrothermal
cycle of erosion, transport, and deposition of sed- venting of Si-charged seawater resulted in rapid
iment operated, but the rivers flowed through a silicification of most near-surface sediments and
landscape that was very different from that of exposed igneous rocks, along with the formation
today. of primary ▶ cherts. Expulsion of fluids at hydro-
The feature that most starkly distinguished the thermal vents led to the deposition of exhalative
Archean and modern land surface was the lack of sediments, variably composed of sulfides, sul-
vegetation. Microbes doubtlessly colonized the fates, carbonates or silica minerals (Russell
subsurface and constructed ▶ biofilms and et al. 2005).
thicker biomats which covered moist areas, pos- The earliest Archean coincided with the end of
sibly including low-lying fluvial floodplains, but the ▶ Late Heavy Bombardment, a time of more
most of the landscape likely had the appearance frequent meteorite impacts. The largest of these
of bare rocks we see on recent Mars. The rate of would have vaporized parts of the oceans, raised
weathering was enhanced by higher temperatures oceanic and atmopsheric temperatures signifi-
and a more aggressive atmospheric composition, cantly, and resurfaced parts of the Earth’s sur-
possibly also by violent thunderstorms, but was face. Whether their overall impact, in particular
slowed down by a lack of humic acid and thick on the biosphere, was local or global, possibly
soils that today greatly enhance weathering rates. extending to sterilization of Earth’s surface envi-
These processes and rates are difficult to quan- ronments, is debated (Abramov and Mojzsis
tify. Erosion was likely enhanced by the lack of 2009).
vegetation and thus lack of stabilized river banks,
but restrained by the overall more modest mean
elevation of continents. Active volcanism cov- See Also
ered much of the surface with lava flows or pyro-
clastic deposits. The Moon, in a closer orbit and ▶ Archean Eon
rotating faster about the Earth, caused higher and ▶ Archean Tectonics
more frequent tides which may have raked the ▶ Barberton Greenstone Belt
Archean Eon 125
▶ Chert
▶ Continental Crust Archean Eon
▶ Craton A
▶ Earth, Formation and Early Evolution Hervé Martin1 and Daniele L. Pinti2
▶ Hydrothermal Environments 1
Laboratoire Magmas et Volcans, Université
▶ Isua Supracrustal Belt Blaise Pascal, OPGC, CNRS, IRD, Clermont-
▶ Komatiite Ferrand, France
▶ Late Heavy Bombardment 2
GEOTOP Research Center for Geochemistry
▶ Ocean, Chemical Evolution of and Geodynamics, Université du Québec à
▶ Oxygen Isotopes Montréal, Montréal, QC, Canada
▶ Pilbara Craton
▶ Silicon Isotopes
▶ Snowball Earth Keywords
▶ Weathering
Continental crust; Greenstone belts; Komatiite;
Plate tectonics; TTG

Synonyms
Abramov O, Mojzsis SJ (2009) Microbial habitability of
the Hadean Earth during the late heavy bombardment.
Nature 459:419–422
Precambrian
Arndt NT (1998) Why was flood volcanism on submerged
continental platforms so common in the Precambrian?
Precambrian Res 97:155–164 Definition
Benn K, Mareschal J-C, Condie KC (eds) (2006) Archean
geodynamics and environments. Geophysical
monograph series. American Geophysical Union The Archean (Archaean in British English) is the
164, p 320 second major period in geological history. Pre-
Eriksson PG (1999) Sea level changes and the continental ceded by the ▶ Hadean and followed by the Pro-
freeboard concept: general principles and application
to the Precambrian. Precambrian Res 97:143–154
terozoic, its start is usually taken as the age of the
Hoffman PF, Kaufman AJ, Halverson GP, Schrag DP oldest preserved rocks, either the 4.0 Ga-old
(1998) A Neoproterozoic snowball Earth. Science (Ga = 109 years = billion years) ▶ Acasta
281:1342–1346 gneisses (Canada) or the 3.85–3.80 Ga-old
Knauth LP, Lowe DR (2003) High Archean climatic tem-
perature inferred from oxygen isotope geochemistry of
Amitsôq gneisses (▶ Greenland). The Interna-
cherts in the 3.5 Ga Swaziland Supergroup, South tional Commission on Stratigraphy (ICS, Jan.
Africa. Geol Soc Am Bull 115:566–580 2013) sets its beginning at the Global Standard
Nisbet EG, Cheadle MJ, Arndt NT, Bickle MJ Stratigraphic Age of 4.0 Ga. The transition to the
(1993) Constraining the potential temperature of the
Archaean mantle: a review of the evidence from
Proterozoic is typically taken at 2.5 Ga, which
komatiites. Lithos 30:291–307 was thought to mark a major change in the
Russell MJ, Hall AJ, Boyce AJ, Fallick AE (2005) On Earth’s geodynamic style and corresponds
hydrothermal convection systems and the emergence roughly to the ▶ Great Oxygenation Event. The
of life. Econ Geol 100:419–438
Sleep NH, Windley BF (1982) Archaean plate tectonics:
Archean thus encompasses an approximately 1.5-
constraints and inferences. J Geol 90:363–379 Ga-long period during which the oldest well-
Young G, von Brunn V, Gold D, Minter WEL preserved rocks formed and life likely originated.
(1998) Earth’s oldest reported glaciation: Its tectonic style was different from today, with
physical and chemical evidence from the Archean
Mozaan Group (2.9 Ga) of South Africa. J Geol
more abundant mantle plumes, greatly
106:523–538 fragmented tectonic plates, and longer
126 Archean Eon
mid-oceanic ridges. It is commonly believed that Hebrides with a debated age of

▶ plate tectonics started in this period. 3,520 160 Ma)
• In Asia, the Siberian Aldan ▶ Shield
(3.5–3.0 Ga), the Indian Dharwar
Overview (3.6–2.5 Ga), and the Sino-Korean cratons
(3.8–3.0 Ga)
Geographical and Temporal Distribution of • In Australia, the Pilbara (3.6–2.5 Ga), Yilgarn
Archean Terranes (2.94–2.63 Ga), Gawler (2.5 Ga), and North-
The Archean eon is characterized by the extrac- ern Australia cratons
tion from the mantle and the subsequent differ- • In Antarctica, the Napier complex
entiation of significant amounts of ▶ continental (orthogneisses dated at 3.95–2.46 Ga)
crust. Indeed, at the end of the Archean eon, • In Africa, the Kaapvaal (3.6–2.5 Ga), Zimba-
probably about 75 % of the juvenile continental bwe (3.5–2.5 Ga), and Madagascar
crust had formed. Large parts of this Archean cratons, as well as the Central and West Africa
crust, named ▶ cratons or shields, have been pre- cratons
served on all continents (Condie 1994; Fig. 1), • In South America, the São Francisco and
including: Amazonian cratons (3.5–2.4 Ga) in Brazil
and the 3.4 Ga Guyana Shield
• In Europe, the 3.1–2.5 Ga Baltic (sometimes • In North America, the Wyoming Province,
referred to as Fennoscandian) and 3.8–3.2 Ga USA (3.5–2.5 Ga); the Superior Province
Ukrainian shields as well as a few outcrops in (3.7–2.7 Ga); the Slave Province (dominated
northern Scotland (Lewisian gneisses in the by 2.73–2.63 Ga greenstone sequences but
180° 120° 60° 0° 60° 120° 180°
60° 60°
40° 40°
20° 20°
0° 0°
20° 20°
40° 40°
60° 60°
180° 120° 60° 0° 60° 120° 180°

Archean terranes Outcropping Covered by sedimentary rocks
Archean Eon, Fig. 1 Geographical distribution of the West Africa Craton, 16 São Francisco and Amazonian
Archean provinces (After Condie 1994; redrawn). 1 Baltic Cratons, 17 Guyana Shield, 18 Wyoming Province, 19
Shield, 2 Ukrainian Shield, 3 Scotland Shield, 4 Siberian Superior Province, 20 Slave Province, 21 Labrador Shield,
Shield, 5 Indian Craton, 6 Sino-Korean Craton, 7 Pilbara and 22 Greenland Shield. Dotted areas represent exposed
craton, 8 Yilgarn craton, 9 Northern Australia craton, 10 Archean terranes, while striped areas represent regions
Napier complex, 11 Kaapvaal craton, 12 Zimbabwe Cra- underlain by Archean rocks
ton, 13 Madagascar Craton, 14 Central Africa Craton, 15
Archean Eon 127
with the ▶ Acasta gneisses dated to 4.03 Ga);

the Labrador Shield (Canada) and the
▶ Greenland Shield (3.8–2.6 Ga with older A
units at ▶ Akilia and Isua up to 3.88 Ga)
The largest continuously exposed outcrop of

Archean rocks is the Amitsôq gneiss in Green-
land with an area of 3,000 km2. The protolith of
these rocks consists of older granitoids, metamor-
phosed into gneisses with emplacement ages of
3.822 0.005 Ga. The oldest supracrustal rocks
(volcaniclastic and sedimentary) are in Akilia
island and the Isua Supracrustal belt with older
ages at 3.872 0.010 Ga, together with banded
iron formation of the Nuvvuagittuq greenstone
belt (3.817 0.016 Ga or older). The recognized
oldest rocks on the Earth (covering a surface of
about 20 km2) are the ▶ Acasta gneisses in Can-
ada (Slave Province) with an age of
4.030 0.003 Ga (Bowring and Williams
1999), while the oldest known minerals are the
famous Jack Hills detrital zircons (Western Aus-
tralia) with recorded ages as old as 4.404 Ga Archean Eon, Fig. 2 Photo of typical gray gneisses
(Wilde et al. 2001). Similarly, inherited cores in (TTG) from Sand River, Limpopo, South Africa. These
zircons from Acasta provided an age of 3.283 0.008 Ga-old rocks consist in the alternation of
whitish quartz-plagioclase layers with biotite and
4.20 0.06 Ga (Isuka et al. 2006). These zircon amphibole-richer gray layers (Photo H. Martin)
crystals are thus the only records of Hadean crust
existing on Earth. Recently, a model age of
4.28 Ga has been proposed for an amphibolitic amphibole-rich layers (Fig. 2). Contrary to typi-
rock (Faux amphibolite), outcropping in the cal modern granites, the TTG contain very low
Nuvvuagittuq greenstone belt (O’Neil proportion of potassic feldspar (KAlSi3O8).
et al. 2008), though this age is strongly debated. The parent magma from which TTG derived
results from the melting at high pressure of a
Geology of Archean Terranes hydrated mafic rock of basaltic composition
Archean terranes all show similar lithological (Fig. 3a). Indeed, when the pressure increases,
associations, independent of their age: (1) granite- basalt is transformed into amphibolite
gneiss, (2) ▶ greenstone belts, and (3) late gran- (amphibole garnet plagioclase feldspar-
itoids. In addition, some show a cratonal cover. rich rock) and then into eclogite
The granitic gneisses are the most abundant, (pyroxene + garnet rock). These products are
composing up to 80 % of the Archean continental melted to give the parental magmas of TTG
crust. Better known under the acronym TTG for (e.g., Martin and Moyen 2002). Although all
▶ Tonalite-Trondhjemite-Granodiorite associa- geologists agree on the basaltic source of TTG,
tion (Jahn et al. 1981), these rocks are coarse- the geodynamic environment where melting took
grained, gray orthogneisses (which means place is still debated: (1) ▶ basalts from a
derived from magmatic rocks, in this case gran- subducted oceanic crust (Fig. 3; Martin 1995;
itoids) with well-developed banding consisting in Martin and Moyen 2002); or (2) underplated
the alternation of whitish quartz-plagioclase basalts melted during the passage of a mantle
layers with gray to black biotite- and plume (Smithies 2000). The first hypothesis can
128 Archean Eon
b
a 40
0 CO cc
5% water 50 ms
Oceanic crust solidus 100
30
dehydratation 100
reaction
Depth (km)
80
P (kbar)
150 Archean
20
60 c
ay
G
d
0 CO
To
s
cc
idu
n 40
ea
sol
10
Arch H 50 ms
Dry
20
100
0 0 Today
0 200 400 600 800 1000 1200 150
T (°c)
Archean Eon, Fig. 3 (a) Pressure-temperature (P-T) Benioff plane was very high; the subducted slab melted
diagram and schematic cross section of both Archean (b) at shallow depth before dehydration could take place.
and modern (c) subduction zones. (a) The P-T diagram After 2.5 Ga, Earth was cooler and the geothermal gradi-
shows the dry and 5 % hydrous solidus of tholeiite as well ent along the Benioff plane was lower (inset c) such that
as main dehydration reactions of oceanic lithosphere. slab dehydration generally occurred before its melting
Abbreviations: H is hornblende out, A is anthophyllite began. The liberated volatiles (mainly water) ascended
out, C is chlorite out, Ta is talc out, Tr is tremolite out, through the mantle wedge, thus lowering its solidus tem-
Z is zoisite out. G outlines stability field of garnet. The perature, which induced its melting. Abbreviations: OC is
grey field is the P-T domain where slab melts can coexist oceanic crust, CC is continental crust, MS is solidus of
with hornblende- and garnet-bearing residue. In the hydrated mantle, black teardrops indicate magma
Archean (inset b), the geothermal gradient along the
be explained if we assume a hotter Archean man- or not to the subduction of a younger oceanic
tle, as inferred by the occurrence in Archean crust, then the latter cannot rapidly dehydrate
times of Mg-rich magmas called ▶ komatiites. and consequently, direct melting of hydrated
It can also be explained by the subduction of basalts is possible, resulting in TTG magma gen-
young and consequently hotter oceanic plates. esis (Fig. 3c).
In modern tectonic regimes, the subducted plate ▶ Greenstone belts represent only 5–10 % of
is old and cold, such that it dehydrates during its the Archean terrains. These elongated structures
descent into the mantle; indeed, there, the oceanic (typically >100 20 km) contain variable
plate is subjected to both higher pressure and amounts of metamorphosed mafic to ultramafic
temperature, such that it dehydrates. Volatiles, volcanic sequences associated with sedimentary
mainly water, are liberated and ascend through rocks. The name “greenstone” comes from the
the mantle wedge, thus lowering its melting tem- green hue imparted by the color of the metamor-
perature (the mantle wedge lies between the phic minerals within the mafic rocks. Chlorite,
descending or subducting oceanic plate and the actinolite, and other green amphiboles are the
continental (or oceanic) plate). Mantle wedge usual green minerals. In some cases, greenstone
melting generates magmas with andesitic to gra- belts show a specific stratigraphic sequence, with
nitic composition that are accreted to form new ultramafic lavas (komatiites) at the base of the
continental crust (Fig. 3b). Modern subduction sequence, followed by basalts (often erupted
systems (both mantle and oceanic crust) are nor- subaqueously with typical pillow structures).
mally too cold to allow subducted basalt melting. Variably metamorphosed sedimentary rocks
If we assume a hotter Archean mantle associated (▶ metasediments) are emplaced at the top of
Archean Eon 129
Archean Eon,
Fig. 4 Pillow lavas of
2.65 Ga-old tholeiitic basalt
from Kuhmo (Finland) A
(Photo H. Martin)
Archean Eon,
Fig. 5 Clastic sediments
(conglomerate) from the
base of the 3.22 Ga-old
Moodies group, Barberton,
South Africa (Photo
H. Martin)
the sequence. Greenstone belts are in complex demonstrate that the internal Earth heat produc-
contact relationships with adjacent plutons or tion in the Archean was higher than today. After
metamorphic rocks: Commonly, the contacts are emplacement, these magmas cooled very rapidly,
structurally modified (thinned or thickened) resulting in acicular and dendritic textures,
contactmetamorphic aureoles; brittle-ductile referred to as “spinifex” textures, which are typ-
fault planes are also common. ical of komatiites. Earth almost totally ceased to
▶ Komatiites are ultramafic volcanic rocks produce komatiites after 2.5 Ga. Archean mafic
(Arndt et al. 2008), almost exclusively restricted volcanics are mainly tholeiitic basalts (Fig. 4),
to the Archean eon, which are distinguished from while calc-alkaline lavas are rare. In contrast to
the more common basalts by a higher content of modern Earth, Archean andesites are rare. At the
MgO (>18 %) and correlated low contents of top of the sequence, more felsic rocks (dacites to
most other elements. The high Mg content is rhyolites) can occur interbedded within the sedi-
explained by a higher degree of melting of the mentary successions.
mantle; the emplacement temperatures of Sedimentary successions include thick
komatiites ranges between 1,400 C and litharenites deposited in deep water as turbidites
1,650 C (Arndt et al. 2008) compared to (Fig. 5) and subordinates conglomerates. Well-
1,100–1,300 C for modern basalts. They preserved 3.5 Ga old turbidites can be observed in
130 Archean Eon
Archean Eon,
Fig. 6 Banded Iron
Formation (BIF) from
Copping Gap (Australia).
These rocks consist of
alternating silica- and iron-
rich layers (Photo
H. Martin)
Archean Eon,
Fig. 7 Dyke of high-Mg
granodiorite (sanukitoid)
intrusive into the 2.65 Ga-
old Kuhmo greenstone belt
(Finland) (Photo H. Martin)
the Komati River valley of the Barberton Green- was modified by assimilation of TTG (Martin
stone Belt, South Africa. They are overlain by et al. 2009).
and interbedded with siltstones, shales, chert, and
banded iron formations (BIF; Fig. 6). Cherts and Archean Geodynamics
BIF are common lithologies in Archean green- Modern plate tectonics induces horizontal forces
stone belts and are likely the result of intense that cause thrusting during orogenesis. These
hydrothermal activity on the ocean floor structures are known in most Archean terranes,
(Westall 2005; Van Kranendonk 2006). indicating that tectonics similar to modern plate
Both the TTG basement and the greenstone tectonics was operating since at least 4.0 Ga ago.
belts were later intruded by high-magnesium However, Archean terranes also show large evi-
granitoids or sanukitoids (Fig. 7). These calc- dence of major vertical deformation that produce
alkaline granites are rich in potassic feldspars dome-and-basin structures (Fig. 8) which are
and magnesium and they might derive by melting exclusive to Archean times. This type of tectonics
of a mantle peridotite, whose initial composition is driven by gravity (as opposed to plate tectonics
Archean Eon 131
Archean Eon, Fig. 8 (Left) Sketch depicting the three creating a sedimentation basin at the center of the green-
main steps of sagduction: (1) In a greenstone belt, high- stone belt. Dark gray is komatiites, light gray is TTG
density komatiites (d = 3.3 g/cm3) emplace over lower basement, and white are sediments. (Right) Satellite
density (d = 2.7 g/cm3) TTG basement rocks, thus gen- photo of the sagduction structures at the Pilbara craton,
erating an inverse density gradient; (2) komatiites sink Western Australia. The greenstone belts (in dark gray) are
downward into the TTG basement which favor a relative localized between TTG domes (white). The width of the
upward motion of the TTG; (3) the movement is amplified photo is 300 km
which is driven by mantle convection) and this of heat to evacuate was greater, it can be con-
process has been known as sagduction since the cluded that the excess of heat has been released
1970s (Gorman et al. 1978). Sagduction struc- through convective processes. Convection rate
tures result from the down motion of high could have been slightly greater, but mainly the
density greenstones (such as komatiites; ridge length was significantly greater than today.
density = 3.3 g/cm3) into the TTG basement The amount of heat dissipated is correlated with
(density = 2.7 g/cm3) and the concomitant cubic square of the ridge length (Hargraves
upward motion of low-density TTG into the 1986). Because the Earth volume and surface
greenstones creating inverse diapirs. At the top did not significantly change since 4.5 Ga, a
of the inverted diapirs, a basin is created allowing greater ridge length should result in smaller
deposition of sedimentary rocks. Several author- plates (Fig. 9). The greater ridge length can also
ities have suggested that horizontal forces acted account for the abundance of cherts and BIFs in
mainly at the plate boundaries (as today) while Archean greenstone belts.
sagduction processes were concentrated within
plates. Hydrosphere, Atmosphere, and Climate
Another difference compared with today is the There is good evidence that oceans were present
supposed length of the mid-ocean ridges, i.e., the on the Earth already in the Early Archean
divergent boundaries between two plates. Indeed, (3.8 Ga ago) after the ▶ Late Heavy Bombard-
the large amounts of internal heat produced dur- ment (Abe 1993; Sleep et al. 2001). The convinc-
ing Archean times has necessarily been released, ing evidence comes from one of the oldest areas
otherwise, the accumulated heat should have of volcanic and sedimentary rocks – the ▶ Isua
resulted in melting the external part of our planet, Supracrustal Belt, West Greenland. The ages of
which is not attested by geological record. As the rocks have been established at about
conduction is not efficient at all to evacuate inter- 3.7–3.8 Ga (for Nuvvuagittuq a date of 4.28 Ga
nal heat, Archean convection should have played has been proposed; O’Neil et al. 2008). In the
this role and, as today, heat must have been Isua Supracrustal Belt, pillow basalts provide
released by ocean-ridge systems. As the amount evidence of underwater eruption and
132 Archean Eon
Archean Eon, Fig. 9 Sketch representing the size of the tectonic plates that presently cover the surface of the Earth
(left) and that supposed for the Archean plates (right)
metasedimentary rocks (banded iron formations, 1 ppmv against the 21 % by volume today.
metapelite, and ferruginous quartzite) are the Oxygen concentrations rose only at the end of
products of erosion, fluvial transport, and sub- Archean to values close to 1 % of their present-
aqueous deposition (Rosing et al. 1996). day level, probably because of shifts in the com-
Primary fluid inclusions were found in quartz petition between the production of oxygen
crystals in iron oxide structures from the 3.5 to derived from cyanobacteria photosynthesis and
3.2 Ga ▶ Barberton greenstone belt, South Africa the rate of consumption of oxygen by different
(Channer et al. 1997), intra-pillow quartz from geological processes. The amount of CO2 in the
the Dresser Formation (3.49 Ga), ▶ Pilbara cra- atmosphere is still a matter of debate, but the
ton, Western Australia (Foriel et al. 2004), and in occurrence of larger amounts of greenhouse
the 2.7 Ga Abitibi Greenstone Belt, Ontario, Can- gases (CO2, CH4) may have been needed in the
ada (Weiershauser and Spooner 2005). The anal- Archean atmosphere to counterbalance the lower
ysis of major cations and anions indicates that the radiation from the faint young Sun, which was
chemistry of the seawater was similar to today, 30 % less than the present-day value. Archean
with some noticeable differences in iron, iodine, terranes do not contain evidence for major glaci-
and bromine abundances indicating a larger influ- ations during the first two billion years of the
ence of hydrothermal fluids (today, the chemistry Earth’s history indicating that a warmer climate
of seawater is mainly controlled by weathering of (as suggested by high ocean temperatures; Robert
continents with a minor role for hydrothermal and Chaussidon 2006) dominated during the eon.
fluids). Salinity was basically NaCl-dominated,
though salinities up to ten times the present Life
values have been measured, possibly related to Though the timing of the origin of life is
seawater evaporation in closed basins (Foriel unknown, the Archean world likely saw the
et al. 2004). emergence of the first organisms. Several mor-
The atmosphere was possibly mildly reducing. phological, molecular, and chemical traces of life
The amount of N2 was likely close to the present punctuate the Archean sedimentary record. Cur-
level (Kasting 1993); CO2 might have been pre- rently, it is difficult to declare with certainty what
sent in larger amounts (up to 1 % in volume or the oldest trace of life is, and importantly what its
higher; Kasting 1987), while oxygen was likely nature and habitat were. Life can be traced
Archean Eon 133
Archean Eon,
Fig. 10 3.5 Ga-old
stromatolites from North
Pole (Pilbara, Australia) A
unambiguously to approximately 2.7–3.0 Ga ago ▶ Craton

(Lopez-Garcia et al. 2006). Beyond this point ▶ Earth, Formation and Early Evolution
many claims for biological processes have been ▶ Earth’s Atmosphere, History of the Origins
made, and all of them have to some degree been ▶ Granite
questioned. Some of the intriguing but controver- ▶ Greenstone Belts
sial early Archean traces include (1) isotopically ▶ Igneous Rock
light graphite inclusions in rocks older than ▶ Isua Supracrustal Belt
3.8 Ga from Akilia island and the Isua ▶ Metamorphic Rock
Supracrustal Belt in southwest Greenland ▶ Metamorphism
(Mojzsis et al. 1996; Van Zuilen et al. 2002); ▶ Metasediments
(2) kerogenous microstructures, stromatolites, ▶ Pilbara Craton
and diverse stable isotope ratio anomalies in ▶ Pillow Lava
3.5 Ga cherts from the Pilbara Granitoid- ▶ Shield
Greenstone Belt in Western Australia (Schopf ▶ Volcaniclastic Sediment
1993; Brasier et al. 2002; Ueno et al. 2006; Pinti
et al. 2009) (Fig. 10); (3) kerogenous microstruc-
tures, stromatolites, and diverse stable isotope References and Further Reading
ratio anomalies in cherts, as well as microscopic
tubes in altered pillow basalts from the 3.4 to Abe Y (1993) Physical state of the very early Earth. Lithos
30:223–235
3.2 Ga Barberton Greenstone Belt in South
Arndt N, Lesher MC, Barnes SJ (2008) Komatiite.
Africa (Staudigel et al. 2008). Cambridge University Press, New York, 488 pp
Bowring SA, Williams IS (1999) Priscoan (4.00–4.03 Ga)
See Also orthogneisses from northwestern Canada. Contrib
Mineral Petrol 134:3–16
▶ Acasta Gneiss
Brasier M, Green O, Lindsay J, Mcloughlin N, Steele A,
▶ Akilia Stoakes C (2005) Critical testing of Earth’s oldest
▶ Amphibolite Facies putative fossil assemblage from the 3.5 Ga Apex
▶ Archean Traces of Life chert, Chinaman Creek, Western Australia. Precam-
brian Res 140(1–2):55–102
▶ Barberton Greenstone Belt
Channer DMDR, de Ronde CEJ, Spooner ETC (1997) The
▶ Basalt Cl-Br-I composition of 3.23 Ga modified seawater:
▶ Canadian Precambrian Shield implications for the geological evolution of
134 Archean Mantle
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dam, 528 pp granodiorite (TTG) series is not an analogue of Ceno-
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B (2004) Biological control of Cl/Br and low sulfate Staudigel H, Furnes H, Mcloughlin N, Banerjee N,
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Rosing MT, Rose NM, Bridgwater D, Thomsen HS Definition
(1996) Earliest part of Earth’s stratigraphic record: a
reappraisal of the >3.7 Ga Isua (Greenland) Archean mantle refers to the terrestrial mantle
supracrustal sequence. Geology 24:43–46 during the Archean eon, which differed both
Schopf JW (1993) Microfossils of the early Archean apex
chert: new evidence of the antiquity of life. Science physically and chemically from the modern-day
260(5108):640–646 mantle.
Archean Tectonics 135
Overview areas of West Greenland have been interpreted to

indicate derivation from strongly depleted man-
The mantle is that part of Earth or other planets tle. Either these rocks were extracted from a small A
between the ▶ crust and the core. The upper and localized volume of mantle or a large volume
mantle, from the base of the crust at about 9 km of continental crust had formed at this time.
(oceanic) or 30 km (continental) to the transition If the mantle were significantly hotter, it
zone at 660 km, is composed mainly of ▶ peri- would have been drier because the reactions that
dotite, an ultramafic rock mainly composed of liberate water in upwelling mantle, where
▶ olivine, pyroxene, and minor garnet. In the degassing takes place, or in subduction zones,
lower mantle, which extends to the core at where the mantle is rehydrated, are temperature-
2,990 km, the minerals are mainly Mg- and dependent. The proportion of water on the sur-
Ca-perovskite ((Mg, Ca)TiO3) and face was larger and therefore the volume of the
magnesiow€ ustite ((Mg, Fe)O). The mantle is oceans may have been greater. The oxidation
solid except for localized zones of partial melt- state of the Archean mantle has been investigated
ing, but it convects with velocities of a few tens of using redox-sensitive elements such as vana-
centimeters per year. dium; no significant difference from that of the
The Archean mantle differed from the modern modern mantle has been established.
mantle in several important ways. Because the
main sources of heat – ▶ radioactivity, residual
heat from accretion, and core
See Also
crystallization – were more active than today,
the mantle was hotter and it convected more
▶ Archean Environmental Conditions
vigorously. Higher temperatures in mantle
▶ Archean Eon
upwelling beneath ▶ mid-ocean ridges produced
▶ Archean Tectonics
larger melt volumes and a thicker oceanic crust.
Higher temperatures at depth may have resulted
▶ Jack Hills (Yilgarn Craton, Western Australia)
in larger and hotter mantle plumes. The abun-
▶ Komatiite
dance of ▶ komatiite only in the Archean pro-
▶ Mantle, Oxidation of
vides evidence of higher mantle temperatures.
▶ Peridotite
The composition may also have been differ-
▶ Ultramafic Rocks
ent, if, as many authors believe, the continental
crust was less voluminous through the Archean.
Continental crust contains far higher concentra-
tions of elements such as Si, Al, K, and the
“incompatible” trace elements, and the segrega- Archean Tectonics
tion of this crust has left the upper part of the
modern mantle depleted in these elements. If Martin J. Van Kranendonk
crustal growth were incomplete in the Archean, School of Biological, Earth and Environmental
either the volume of depleted mantle or the Sciences, University of New South Wales,
degree of depletion would have been less. The Australia
isotopic composition of rocks from the Archean
mantle should, in theory, cast some light on the
problem but at present the message is ambiguous. Keywords
The Hf isotope compositions of ▶ zircons show
evidence of extraction prior to 4 Ga ago of Archean; Continents; Crust; Lithosphere; Non-
enriched material, perhaps continental crust; the uniformitarian; Plate tectonics; Tectonics; Ther-
Nd isotopic compositions of rocks from the oldest mal evolution of the Earth; TTG
136 Archean Tectonics
Synonyms steeply dipping, generally synclinal, greenstone

keels (granite-greenstone crust: Hickman 1984;
Crustal deformation; Early earth Chardon et al. 1996); and a unique type of ultra-
mafic lava known as ▶ komatiite derived from
high-temperature mantle melts (Viljoen and
Definition Viljoen 1969; Arndt 2003). Many authors also
noted that Archean granite-greenstone crust
Archean tectonics is the study of the formation, lacked the diagnostic features of modern subduc-
interaction, and deformation of the Earth’s conti- tion/collision zones, including accretionary tec-
nental and oceanic crust during early Earth his- tonic mélange, ophiolites, and high-pressure/
tory (the Archean Eon; ca. 4.0–2.5 Ga) and the low-temperature metamorphism, leading some
driving forces behind these processes, including to suggest – even recently – that plate tectonics
mantle plumes, subduction, and accretion/colli- either did not operate in the Archean (Hamilton
sion. This topic remains highly controversial due, 1998; McCall 2003; Stern 2005) or operated in
in part, to a fragmentary rock record, but also to conjunction with other processes (e.g., Rey
nonunique interpretations of complex geological et al. 2003; Sandiford et al. 2004).
datasets in the absence of actualistic plate config-
urations. Historically, Archean tectonics has been
polarized into uniformitarian (i.e., analogous Overview
with modern, or Phanerozoic Earth) and
non-uniformitarian views, but recent studies The Archean ▶ mantle was probably 100–300 K
have favored modern Earth processes in the hotter than today, which significantly affected the
Archean, complicated by problems arising dominant tectonic style. Geochemical, geophysi-
mainly from greater heat production and higher cal, and modeling evidence suggests some form
mantle temperatures (Condie 1994; Benn of ▶ plate tectonics in the Archean, although the
et al. 2006; Brown and Rushmer 2006). The dis- absence of key characteristics such as ophiolites
cussion revolves around the basic question if and and blueschists implies that it probably differed
how tectonics in the Archean was different from from modern ▶ plate tectonics. Evidence for
modern-style plate tectonics. Archean plate tectonics was recognized quite
early on in the type of Archean ▶ crust known
as high-grade gneiss terranes. This evidence
History included the presence of large-scale recumbent
isoclinal folds associated with crustal thickening
Although the relative antiquity of some parts of (Bridgwater et al. 1974; Myers 1976; Wilks
the continental crust was recognized more than 1988; Hanmer and Greene 2002), voluminous
150 years ago (Logan 1857), it was not until the sodic granitoids derived from high-pressure melt-
advent of radiometric dating, 100 years later, that ing of basalt (Martin et al. 2005;
the antiquity of much of the continental crust was Rapp et al. 1991), high-pressure metamorphism
fully appreciated (Stockwell 1961). Only in the (Riciputi et al. 1990; Harley 2003), and structures
past decade it has been discovered that the pre- consistent with terrane accretion (Nutman
served crustal record on Earth extends back to et al. 2002; Windley and Garde 2009). Over the
within 150 Ma of the age of formation of the past three decades, abundant evidence for
Solar System (Wilde et al. 2001). Early geologi- Archean plate tectonics has also been found in
cal studies found that continental crust older than some Archean granite-greenstone terrains in the
about 2.5 Ga was different from younger crust. form of thrusts and recumbent isoclinal folds,
Archean crust showed distinct regional patterns coupled high-pressure-low-temperature meta-
defined by overlapping, elliptical areas of granitic morphism, fossil subduction zones, accreted ter-
rocks (gregarious batholiths; Macgregor 1951); ranes, rift sequences, and subduction-zone
magmatism (Card 1990; Heubeck and Lowe subduction), blueschists, and ultrahigh-pressure
1994; Calvert et al. 1995; Smithies et al. 2005; metamorphic rocks (partly subducted rocks that
Moyen et al. 2006; Wyman et al. 2006; Van emerged again at the Earth’s surface) are all well- A
Kranendonk 2007; Van Kranendonk understood features associated with modern plate
et al. 2010). However, an absence of hallmark tectonics. Their occurrence throughout Earth’s
characteristics of modern subduction-accretion history is thought to provide clear indicators of
zones in many granite-greenstone terrains plate-tectonic activity. Large-scale tectonic
(Hamilton 1998; McCall 2003; Stern 2005), the structures can be indicative as well: linear struc-
autochthonous nature of some major greenstone tures are often interpreted as remnants of subduc-
successions, and suggestions that mantle roots tion trenches, whereas large oval-shape structures
form through in situ melting events rather than are thought to be diapir- or dome-related that
through subduction stacking indicates local might not need any plate-tectonic activity.
crustal development as volcanic plateaus devel- Geochemical and petrologic techniques pro-
oped on top of older continental basement (e.g., vide the “fingerprints” of the chemical processes
Blenkinsop et al. 1993; Bleeker et al. 1999; Van associated with Archean tectonics. Mantle melt-
Kranendonk et al. 2007). Indeed, many studies ing will deplete the mantle of “incompatible”
suggest that some pieces of Archean crust contain elements (Rollinson 2007). Another form of ele-
features that cannot be ascribed to uniformitarian, ment separation occurs due to differences in fluid
Phanerozoic-type, plate tectonics, but rather mobility, so that some elements will preferen-
formed as a result of large-scale infra-crustal tially move with any pore fluids, while others
differentiation accompanying periods of mantle- will stay in the residue. These processes will
plume-related magmatism (Stein and Hofmann leave geochemical fingerprints that can be used
1994; Whalen et al. 2002; Rey et al. 2003; Smith- to recognize ancient subduction processes (e.g.,
ies et al. 2009; Van Kranendonk et al. 2009). The Shirey et al. 2008). In addition, geochemical dat-
fact that different processes have been recognized ing of crustal rocks and mantle material shows
from studies of different pieces of Archean crust how mantle material was depleted through time
indicates that there was no single Archean tec- by continent formation. Since continents today
tonic process, but rather that – as with modern are formed primarily at subduction zones, this
Earth – Archean continental crust formed through has been interpreted as another indication for
a variety of processes, including plate tectonics the presence of ancient subduction and therefore
and mantle-derived upwellings, and probably plate tectonics.
through the interaction between these two Various modeling techniques are used to fur-
end-member processes. ther constrain the range of dynamically viable
tectonic processes during the Archean. Tectonic
vigor is related to mantle convection and inti-
Basic Methodology mately couples to the cooling rate of the Earth:
tectonic activity leads to increased heat loss from
Direct field evidence from Archean continental the mantle, and mantle temperature influences the
lithosphere provides a record of Archean tectonic vigor of tectonic activity. Mantle temperature
processes. Geophysical methods include paleo- through time therefore provides an important
magnetism and seismic evidence. Solidifying constraint, and parameterized and numerical
magma registers the paleo-latitude and thus can modeling techniques provide a means to link
record (relative) continental motion. Seismic pro- mantle temperature and tectonic activity.
files through Archean crust indicate the presence Today’s plate tectonics is primarily driven by
of dipping seismic reflectors, which could be the subduction process, and subduction dynamics
interpreted as the remnants of a fossil subduction is, to a large extent, influenced by mantle temper-
zone. Among the geological evidence, ophiolites ature due to melting events and the temperature-
(slivers of oceanic lithosphere that escaped dependent strength of the lithosphere.
The viability and style of subduction in an early, flux was higher or that Earth was not cooling
hotter Earth is investigated using parameterized (significantly) or was perhaps even heating
and numerical modeling techniques. up. Inferred liquidus temperatures from
ophiolites and greenstone belts suggest a gradual
mantle temperature drop of 200 K since the
Key Research Findings Archean. Komatiitic melt data suggest a mantle
potential temperature (i.e., mantle T extrapolated
Field evidence has been used to argue for or to surface P, T-conditions) reduction by300 K
against modern-style plate tectonics in the for dry melting to 100 K if melting took place
Archean. Ophiolites are preserved pieces of old under much wetter conditions. Jaupart
oceanic lithosphere that escaped subduction, and et al. (2007) provide a recent overview of the
the occurrence of old ophiolites is therefore a thermal evolution of the Earth.
clear indicator of plate-tectonic activity. They The dynamical viability of subduction in the
are widespread since 1 Ga, but are much rarer Archean has been questioned. Today, plate tec-
before that. Recently, Furnes et al. (2007) tonics is primarily driven by dense slabs sinking
reported a 3.8 Ga-old ophiolite in Isua, West into the mantle, and thereby pulling the trailing
Greenland, although their interpretation has lithosphere across the surface. Archean plate tec-
been disputed. Other direct types of evidence tonics would require a similar driving mecha-
for plate tectonics are blueschists and ultrahigh- nism. A hotter mantle provides more melt and
pressure metamorphic rocks, which are both gen- therefore a thicker, low-density mafic crust (up to
erally believed to form within subduction zones, 20 km thick instead of today’s 5–8 km) that
where they are brought down to large pressures doesn’t easily subduct. Although dehydration
and temperatures, and subsequently make it back during melting will make the plate composition-
to the surface. The oldest blueschists are ally stronger and could allow for similar plate-
ca. 850–700 Ma old, while the oldest UHP local- tectonic rates in the Archean as today (Korenaga
ities are 600 Ma old. These data could indicate 2006), thermal weakening would probably dom-
that modern-style plate tectonics did not start inate if the mantle was substantially hotter and
until the Neoproterozoic (Stern 2005) or that the would result in weaker plates. The combined
appearance of plate tectonics evolved over time buoyancy and plate strength effects make sub-
(van Hunen and van den Berg 2008). duction inefficient for mantle temperatures more
Earth’s thermal evolution provides further than 150 K hotter than today (van Hunen and van
constraints. Today, plate tectonics forms the den Berg 2008).
dominant cooling mechanism for the Earth and Today’s continental crustal rocks (loosely
is therefore closely linked to the thermal evolu- termed andesites) differ significantly in trace-
tion of the Earth. The Archean Earth had an element composition from their Archean coun-
amount of radiogenic heat production two to terparts (trondhjemite-tonalite-granodiorite, or
three times larger than today due to the gradual TTGs). Whereas andesites are thought to ulti-
decay of the dominant heat-producing elements mately derive from melting in the hydrated
uranium, thorium, and potassium in the mantle. supra-subduction mantle wedge, TTGs seem to
Today, the surface geothermal heat flux (heat form from wet melting of oceanic basalts at
escaping the Earth’s interior) is 30 % provided sometimes >50 km depth, and the most popular
by internal radiogenic heating while the formation model is melting of subducting oceanic
remaining 70 % comes from cooling of the crust (Foley et al. 2002). So the differences
Earth (Turcotte and Schubert 2002; Korenaga between modern and Archean continental crust
2006). This shows that surface heat flow from suggest a secular evolution of the subduction
radiogenic heating was a lot more important in process. But at the same time, it indicates the
the Archean which implies that either surface need for a process to bring fluids to 50–100 km
tectonics were such that the total surface heat depth throughout the Earth’s history. At present,
no other mechanism than subduction seems capa- Venus (which doesn’t have plate tectonics, prob-
ble of doing that, which is regarded as one of the ably because of the lack of liquid water, but
strong arguments in favor of Archean plate perhaps experiences episodic large-scale mantle A
tectonics. overturns).
However, although most studies support
Archean plate tectonics, perhaps in some modi-
fied form, the possibility of other dominant tec- Future Directions
tonic processes should not be excluded. If indeed
plate tectonics were absent in the Archean, such Integrated, four-dimensional lithospheric studies
alternative tectonic models were probably essen- of Archean lithosphere are the key future research
tial to provide a mechanism for the observed directions, particularly in poorly studied regions.
steady mantle cooling of 50–100 K/Gyr. One Detailed geochronology within a well-
popular model is the crustal delamination model established map framework continues to be key
(e.g., Zegers and van Keken 2001), in which to understanding formation processes of ancient
mantle melting events would thicken the conti- crust, particularly when coupled with ongoing
nental crust until its base becomes gravitationally reevaluation of uniformitarian assumptions
unstable and would cause lithospheric overturn. given known aspects of secular change. Hf iso-
Such models would explain observations from tope determination of zircon can help discrimi-
early Archean rocks such as the ovoid-shaped nate juvenile crustal growth through subduction
intrusions in the eastern Pilbara craton. from volcanism and crustal recycling during epi-
sodes of plume magmatism. Further understand-
ing of crustal growth processes will be aided by
Applications more complete knowledge on the origin of sub-
continental lithospheric mantle: detailed studies
The style and vigor of tectonics in the Archean of primary dunite-harzburgite xenoliths are
has important consequences for many aspects of required to determine the age, composition, and
the evolution of the Earth. Tectonic style directly history of these more depleted rocks, and their
influences (a) how and when continents formed properties tied into detailed regional seismic
and why cratons remained stable and preserved studies, including physical modeling of their geo-
over much of the Earth’s history (Lenardic physical response. Additional work is also
et al. 2003); (b) how tectonics-related events required on the ▶ metamorphism of granite-
such as melting, remelting, and fluid-related greenstone terranes, specifically precise dating
alteration has changed the composition of the of mineral assemblages related to magmatic and
mantle from its primitive composition shortly deformational events and P-T studies of granitic
after core formation to its modern composition rocks as a counterpart to greenstones, to test
(Shirey et al. 2008); and (c) the composition of models of cold greenstone diapirs in hot rising
the atmosphere and oceans through ▶ degassing granites (partial convective overturn; Smithies
(during volcanism) and regassing (at subduction et al. 2009). Additional studies are required on
zones) of volatiles and surface weathering (Lowe the origin of Archean calc-alkaline felsic volca-
and Tice 2007; Rollinson 2007), and through that nic rocks and high-Mg diorites to establish
the emergence and evolution of life on Earth. whether they are really the products of volcanic
Furthermore, if plate tectonics has been operative arc magmatism over an active subduction zone,
throughout the changing conditions of the Earth as widely assumed, or the products of fraction-
during its history, how does that relate to the ation and crustal contamination of large tholeiitic
viability of plate tectonics on the other terrestrial magma chambers derived from mantle plumes.
planets of our solar system, such as on Mars Additional research is required across the interval
(which has no plate tectonics today, but might 3.3–2.9 Ga, in order to assess the tantalizing
have had some during its earliest history) or clues that there may have been a global change
in crust-formation processes at this time, perhaps images of a mantle suture in the Superior Province.
due to an increase in plate size and concomitant Nature 375:670–674
Card KD (1990) A review of the superior province of the
cooling of oceanic lithosphere and steepening of Canadian shield, a product of Archean accretion. Pre-
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stone terrains in the Archean Dharwar craton (southern
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oceanic changes in the Archean, to the habitat Pilbara Craton, Western Australia: driving mecha-
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Furnes H, de Wit M, Staudigel H, Rosing M,
Muehlenbachs K (2007) A vestige of earth’s oldest
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▶ Degassing Hamilton W (1998) Archean magmatism and deformation
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▶ Komatiite Hanmer S, Greene DC (2002) A modern structural regime
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of Archean granulite metamorphism in the Godthab- Africa, with implications for Archean tectonics. Chem
Fiskenaesset region, southern West Greenland. Geol 261:114–138
J Metamorph Geol 8:171–190 Van Kranendonk MJ, Smithies RH, Hickman AH, Win-
Rollinson H (2007) Early earth systems: a geochemical gate MTD, Bodorkos S (2010) Evidence for
approach. Blackwell, Maldon Mesoarchean (3.2 Ga) rifting of the Pilbara Craton:
Sandiford M, Van Kranendonk MJ, Bodorkos S (2004) the missing link in an early Precambrian Wilson cycle.
Conductive incubation and the origin of dome-and- Precambrian Res 177:145–161
keel structure in Archean granite-greenstone terrains: Viljoen MJ, Viljoen RP (1969) The geology and geochem-
a model based on the eastern Pilbara Craton, Western istry of the lower ultramafic unit of the Onverwacht
Australia. Tectonics 23:TC1009. doi:10.1029/ Group and a proposed new class of igneous rocks. Geol
2002TC001452 Soc S Afr Spec Publ 2:55–86
Shirey SB, Kamber BS, Whitehouse MJ, Mueller PA, Whalen JB, Percival JA, McNicoll VJ, Longstaffe FJ
Basu AR (2008) A review of the isotopic and trace (2002) A mainly crustal origin for tonalitic granitoid
element evidence for mantle and crustal processes rocks, Superior Province, Canada: implications for late
in the Hadean and Archean: implications for the Archean tectonomagmatic processes. J Petrol
onset of plate tectonic subduction. In: Condie KC, 43:1551–1570
142 Archean Traces of Life
Wilde SA, Valley JW, Peck WH, Graham CM (2001) Evi- S especially. These different biosignatures are
dence from detrital zircons for the existence of conti- yet to provide a consistent and complete picture
nental crust and oceans on the Earth 4.4 Gyr ago.
Nature 409:175–178 of early Archean ecosystems, and there is cur-
Wilks ME (1988) The Himalayas – a modern analogue for rently little scientific consensus about when and
Archean crustal evolution. Earth Planet Sci Lett where life first emerged on Earth. Refining our
87:127–136 understanding of microbial biosignatures in the
Windley BF, Garde AA (2009) Arc-generated blocks with
crustal sections in the North Atlantic craton of West Archean rock record is essential to designing
Greenland: crustal growth in the Archean with modern strategies for seeking life elsewhere in our uni-
analogues. Earth Sci Rev 93:1–30 verse and for ratifying this evidence.
Wyman DA, Ayer JA, Conceição RV, Sage RP
(2006) Mantle processes in an Archean orogen: evi-
dence from 2.67 Ga diamond-bearing lamprophyres
and xenoliths. Lithos 89:300–332 Overview
Zegers TE, van Keken PE (2001) Middle Archean conti-
nent formation by crustal delamination. Geology This entry first explains where to look for
29:1083–1086
Archean traces of life, what evidence astrobiolo-
gists seek, and how these rocks are investigated,
especially the techniques and approaches
involved. Then a review of current research find-
Archean Traces of Life ings is given focusing on selected case studies of
microfossils, stromatolites, and microborings
Nicola McLoughlin along with stable isotope evidence from the
Department of Earth Science and Centre for early Archean. Lastly, new frontiers in early
Geobiology, University of Bergen, Bergen, earth research are described with the aim of better
Norway understanding the nature of Archean life and
environments and how this translates to recogniz-
ing signatures of life beyond earth.
Keywords
Earliest evidence of life on earth; Emergence of Basic Methodology

life; Oldest fossils
Locating Well-Preserved Archean Rocks from
Habitable Environments
Synonyms The search for Archean traces on life relies upon
geological mapping and radiometric dating to
Archean biosignatures; Trace of life locate rocks of Archean age. Worldwide, there
are two ▶ cratons that preserve intact sequences
of early Archean age where now metamorphosed
Definition volcanic and sedimentary rocks are preserved in
▶ greenstone belts – so called because of the
The ▶ Archean is the period of geological time green color conferred by their typical metamor-
between 3.8 and 2.5 billion years ago when life is phic minerals. The first of these is the Pilbara of
thought to have emerged on Earth. Traces of Western Australia and the second is the Kaapvaal
Archean life are preserved in rare, fragmentary, Craton of South Africa and the ▶ Barberton
and often highly altered rock sequences. Morpho- Greenstone Belt. In recent years, there have
logical evidence for Archean life is provided by been several Archean scientific drilling projects
▶ microfossils, microborings, ▶ stromatolites, that have targeted these sequences to seek the
and wrinkle mats. Chemical evidence for life is earliest evidence for life. Scientific drilling yields
recorded by stable isotope ratios of C and continuous sequences of rock unaffected by
Archean Traces of Life 143
alteration at the earth’s surface allowing more surface environments were largely anoxic, the
complete investigation of well-preserved atmosphere was probably rich in carbon dioxide
biosignatures within their geological context. and methane, and there was no ozone layer. Sec- A
Older Archean rocks between 3.8 and 3.7 Ga ondly, seawater was supersaturated with silica,
from the ▶ Isua supracrustal belt of Western and its pH, temperature, and salinity are widely
Greenland and also Labrador are of much higher debated and certainly differed from today, with
metamorphic grade and more intensely the ocean temperatures likely being higher.
deformed. Thus, any morphological traces of Thirdly, exactly when Archean ▶ plate tectonics
life have been almost completely destroyed in began, the nature of early tectonic processes is
these earliest rocks, and discussions regarding unclear with profound implications for the
the evidence for life center on chemical evidence cycling of nutrients on Earth and the oxidation
alone. of key geological reservoirs. A comprehensive
The search for Archean life has traditionally review of the latest research investigating early
centered on metasedimentary rocks, in particular Archean rocks can be found in Van Kranendonk
cherts and carbonates. Geological environments et al. (2007).
where microbial remains are most likely to be
preserved are those where rapid, contemporane- Textural Evidence of Life: Investigating
ous mineralization entombs and permineralizes Morphological Complexity
living organisms. Precipitation of microcrystal- Morphological ▶ complexity is often regarded as
line silica is a good example and can preserve a diagnostic criterion for life. But it must be
cellular remains with exceptionally high fidelity. remembered that complex shapes do not require
▶ Cherts are formed in the vicinity of hydrother- complex causes and can arise naturally in
mal vents, hot springs, and as chemical sediments physiochemical systems as shown, for example,
on the Archean seafloor. The retention of fossil- by snowflakes growth. There are three key mor-
ized biosignatures over geological time frames is phological traces of life: ▶ microfossils, ▶ stro-
increased if the host rock comprises phases that matolites, and microborings, and interpretation of
are resistant to postdepositional alteration pro- this morphological evidence proceeds hand in
cesses such as diagenetic recrystallization, disso- hand with studies of modern analogs and explo-
lution, or replacement. In recent years, a new ration of potential abiological mimics as
approach to seeking traces of Archean life has explained below.
come to prominence, and this involves seeking
“footprints” of life or tunnels created by microbes Chemical Evidence of Life: Elemental and
that etch rocks rather than the organic remains of Isotopic Signatures
the microorganisms themselves. Meta-volcanic ▶ Isotopic ratios preserved in ancient rocks may
glass from Archean seafloor ▶ pillow lavas has record past biological activity and can be mea-
been found to contain such “microbial foot- sured by mass spectrometry to test for the pres-
prints,” and Archean carbonate sequences are ence of life and sometimes identify the
now also being reexamined for evidence of such metabolisms involved. Carbon and sulfur are the
rock-tunneling microorganisms. main isotopic tools used in the search for Archean
An important criterion for establishing the life, and these are introduced below. Elemental
▶ biogenicity of candidate Archean traces of mapping in the vicinity of microbial remains
life is the demonstration that the geological envi- can also be highly informative as the life
ronment was viable for life. This translates to the activity and/or the subsequent decay of a micro-
assessment of habitability or, in other words, organism can modify the composition of the sur-
mapping out the environmental limits to life. rounding minerals. Examples are given below
There are a number of first-order differences along with the techniques and instruments capa-
between the Archean world and recent Earth ble of undertaking such isotopic and elemental
that should be borne in mind. Firstly, Archean mapping.
Carbon Isotopes sulfate reduction (Canfield and Raiswell 1999).

Carbon isotopes are probably the most studied In older rocks such fractionations are much
isotopic tracer of life on Earth. Carbon isotope smaller with most sedimentary sulfides older
systematics is described elsewhere in this vol- than 2.7 Ga showing a narrow d34S range, and
ume, and rather here the focus is on their appli- this has several possible explanations. Firstly, a
cation to seeking traces of Archean life. The d13C nonbiological origin from H2S derived from
of the biosphere through time has been measured hydrothermal or volcanogenic processes always
directly from carbonaceous material found in needs to be tested. Secondly, sulfate reducers
ancient sediments, and if this can be shown to only discriminate sulfur isotopes when seawater
be both syngenetic and endogenetic, then it sulfate concentrations are above 1 millimolar.
records the microbial metabolisms employed at Thus, the absence of a large d34S signal before
that time. Typical Archean organic matter is 2.7 Ga could mean that either seawater sulfate
found to have a d13C value of 20 ‰ relative levels were low or that sulfate reduction had not
to inorganic carbonate leading many researchers yet evolved. Evidence in support of the former
to claim that biological activity began 3.8 billion explanation is discussed below.
years ago (Schidlowski 2001). Specific microbial
metabolisms have also been inferred on the basis Additional Stable Isotope Systems
of the magnitude of carbon isotopic fraction- Additional isotopic systems that have been uti-
ations measured from Archean rocks, and several lized to investigate Archean environments and
examples will be discussed below. traces of life are explained in a rich literature
that includes silicon, oxygen, and ▶ deuterium
Sulfur Isotopes isotopes measured on cherts to investigate sea-
Microorganisms that metabolize sulfur com- water temperatures (e.g., Hren et al. 2009),
pounds are one of the most deeply rooted groups ▶ nitrogen isotopes measured especially on
in the tree of life. ▶ Sulfur isotopes preserved in ▶ kerogen (e.g., Godfrey and Falkowski 2009),
ancient sulfides especially pyrite along with sul- and ▶ iron isotopes in a range of rock types (e.g.,
fate minerals like barite are used to trace ancient Dauphas et al. 2004).
microbial metabolisms. A review of sulfur iso-
topes in early Archean rocks can be found in Van Instrumental Techniques for Seeking Traces
Kranendonk et al. (2007). The baseline for of Archean Life
interpreting such data is provided by studies of Having outlined the morphological and chemical
modern microbes that employ sulfur-based basis for seeking Archean traces of life, the tech-
metabolisms including sulfur oxidation, sulfate niques used for deciphering these traces are
reduction, and sulfur disproportionation reviewed. All such investigations begin with geo-
(Canfield 2001). ▶ Sulfur cycling and isotope logical mapping to identify the nature of the host
systematics are explained in detail elsewhere in rocks, to determine if the context was plausible
this volume. Of particular interest to Archean for life, and to establish age relationships with
studies is the development of mass spectrometry other rock units. The key techniques are:
techniques that can measure mass-independent
sulfur isotope fractionations (MIF) as will be Optical microscopy: Examines the morphology
explained below. Sulfur isotopes have also proof the putative biosignature in two dimensions
vided a hotly debated tracer for the rise of atmo- and if z-plane stacking is available in three
spheric oxygen (e.g., Farquhar et al. 2000), and dimensions and also the mineralogy of the
this will not be discussed further here. enclosing rock and relative age of the candi-
The S isotope record from 2 Ga onward date biosignature with respect to other fabrics
shows d34S fractionations of 50–60 ‰ between in the rock.
sulfides that are depleted relative to coexisting Scanning electron microscopy – with energy-
sulfates, and this has been attributed to microbial dispersive X-rays (SEM-EDX): Examines the
shape and surface morphology of a putative molecules that are ▶ biomarkers for specific
biosignature. Accompanying element distri- groups of organisms, for example,
bution maps can be created using EDX. ▶ cyanobacteria. A
Focused ion beam milling – transmission elec- Radiogenic isotopes: The abundances of natu-
tron microscopy (FIB-TEM): FIB is used to rally occurring radioactive elements are mea-
mill a very thin 100 nm wafer from a chosen sured to calculate absolute ages of rocks.
site within a sample targeting, for example, Synchrotron X-ray tomography: Nondestructive
fossilized cell walls. This can then be imaged 3-D morphological images created from a
by TEM at the nanometer scale to reveal cel- series of 2-D X-ray images taken around a
lular and crystalline structures. Electron dif- single axis of rotation. It yields spectacular
fraction patterns can also be generated to images of paleontological samples.
identify crystalline phases. Synchrotron X-ray spectroscopy and microscopy:
Electron microprobe: Is used for nondestructive Uses the absorption of X-rays to image sam-
analysis of the chemical composition of a ples at the micron to nanometer scale and to
biosignature including the quantification of investigate, for example, the redox state or
elements present at levels as low as 100 ppm. coordination chemistry of the sample. There
Confocal laser Raman microspectroscopy: Gen- are many astrobiological applications, for
erates spectra that are diagnostic of different instance, to investigate microbe-mineral
mineral and organic polymorphs and can be interfaces.
used for rapid mineral identification. Also the
spectra can be used for nondestructive 2-D and
3-D morphological mapping of, for instance, Key Research Findings
microfossils. Raman microspectroscopy is
also useful for thermometry, providing crucial Microfossils
assessment of the thermal maturity of ▶ Microfossils are the permineralized remains of
organics, important to evidence their carbonaceous microbial cells and display a range
▶ syngenicity. of shapes that in the Archean include coccoids or
Atomic force microscopy (AFM): Can be used to simple spheres and filaments that may be septate
image and measure the atomic surface struc- and/or branched. An instructive example of how
ture of a sample at the nanoscale by “feeling” the ▶ biogenicity of candidate Archean microfos-
the surface with a cantilever tip (can be sils is assessed comes from the 3.45 Ga ▶ Apex
coupled to a Raman microscope). Chert of Western Australia that is now famous for
Laser ablation inductively coupled plasma mass engendering a vigorous debate regarding the
spectrometry (LA-ICP-MS) of stable isotopes oldest microfossil-like objects (Fig. 1c). In the
C and S: Can measure isotopic ratios for a 20 years since their discovery, these “microfos-
target spot several microns across to detect sils” have become the cornerstone of textbook
potential biological processing of these descriptions of an early Archean biosphere. This
elements. changed, however, when a reexamination of the
Secondary ion mass spectrometry (▶ SIM S and “microfossils” called into question their
nanoSIMS): A surface analytical technique biogenicity (Brasier et al. 2002). These authors
that enables in situ elemental mapping of argued that the geological context, morphology,
major and trace elements and measurement and distribution of the “microfossils” are more
of isotopic ratios at the micron scale or sub- consistent with an origin as ▶ abiotic graphite
micron scale in the case of nanoSIMS. Can artifacts, produced by the recrystallization of
detect elements present in the parts per billion amorphous silica to spherulitic chert. The princi-
range. pal lines of evidence from Brasier et al. (2005)
Gas chromatography – mass spectrometry are summarized in Table 1 and contrasted with
(▶ GC/MS): Used to identify organic the original interpretation of Schopf and Packer
Archean Traces of Life, Fig. 1 Precambrian morpho- composed of carbonaceous material (orange) in a silica
logical traces of life. (a) Photomicrograph of microbial matrix (yellow) from the 3.45 Ga Apex Chert of West
biotextures in an inter-pillow breccia from the Australia; (d) transmitted light image of coccoid micro-
Hooggenoeg Formation of the Barberton Greenstone fossils from the 1 Ga Boorthana Chert of South Austra-
Belt, South Africa, and the meta-volcanic glass comprises lia; (e) putative microbial mat layer from the 3.5 Ga
a greenschist facies assemblage of chlorite and quartz with Buck Reef Chert of South Africa; a lower layer of
titanite-filled tubular texture, and these are curvilinear and rounded, composite, carbonaceous grains is overlain by
unbranched, radiate from a central “root zone,” and are putative microbial mat rip-up fragments that show plastic
segmented by crosscutting chlorite; (b) scanning electron deformation features; (f) automontaged, transmitted light
micrograph of a twisted filamentous pseudofossil made image of intertwined filamentous microfossils from the
experimentally by precipitating barium-carbonate crystals 3.2 Ga Sulphur Springs Group of West Australia. Scale
in sodium silicate gel; (c) branched, septate “microfossil” bars: (a) 50, (b) 20, (c) 10, (d) 50, (e), and (f) 50 mm
Archean Traces of Life, Table 1 Contrasting lines of evidence and their interpretation collected from the 3.45 Ga
Apex Chert of Western Australia and contained “microfossil” structures
Lines of
evidence Brasier et al. Schopf et al.
A
Environment Deep-marine seafloor cherts with intrusive Shallow-marine silicified sediments
of deposition hydrothermal dike cherts
“Microfossil” Sheets and wisps of carbonaceous material Eleven taxa of filamentous “microfossils”
morphology concentrated around the rims of silica
spherulites and rhombic crystal inclusions
Laser Raman The carbonaceous material has a graphitic The “microfossils” have a Raman signature that
analysis Raman signature, and the “microfossil” is argued to be comparable to disordered
signature is indistinguishable from the matrix kerogenous carbon from younger biogenic
carbonaceous material assemblages
Carbon d13Corg of 30 ‰ to 26 ‰ which cannot d13Corg of 30 ‰ to 23 ‰ lies within the
isotopes exclude abiotic Fischer-Tropsch synthesis range of biological fractionation
Interpretation Abiotic artifacts created by the recrystallization Silica permineralization of filamentous
of amorphous silica that displaced graphitic “microfossil” cells that could have included
margins forming a spectrum of arcuate to oxygen-producing cyanobacteria and possibly
dendritic artifacts larger, beggiatoacean microfossils
(1987) and Schopf (2002). An origin for instructive example of how the biogenicity and
these “microfossils” as oxygen-producing antiquity of microfossil-like structures can be
cyanobacteria-like organisms now seems highly tested.
unlikely. Further examples of putative Archean micro-
There have been several subsequent studies fossils including spheroids, ellipsoids, and fila-
that have investigated the morphology of the ments have been reported from the 2.6 Ga
Apex microfossil-like structures and compared Ghaap Subgroup of South Africa, the 2.7 Ga
them to younger less metamorphosed samples Tumbiana Formation of West Australia, the
and also studies that have looked at the carbon 3 Ga Farrell Quartzite of West Australia,
ultrastructure and likened this to younger biogenic the 3.2 Ga Dixon Island Fm of West Australia,
microfossils. But no studies have yet investigated the 3.41 Ga Kromberg Fm of South Africa,
the correlation between seafloor-hydrothermal and the 3.5 Ga Dresser Formation of the Pilbara;
depositional gradients within the Apex Chert and these are illustrated and discussed in Wacey
changes in the candidate chemical and/or morpho- (2009). The perennial difficulty with interpreting
logical biosignatures to conclusively reconstruct a all such structures is that they comprise shapes that
microbial ecosystem. Moreover, SEM investiga- can be difficult to distinguish from natural abiotic,
tions of the Apex Chert have revealed multiple crystal habits that could grow under similar con-
episodes of hydrothermal alteration at tempera- ditions and develop complex self-organized mor-
tures
250 C and also recent groundwater alter- phologies (Brasier et al. 2006). This has been
ation that have generated micrometer-sized silica illustrated by “crystal-garden”-type experiments
structures resembling microbial ▶ exopolymers that precipitate microfossil-like biomorphs in
and textures formed by the partial dissolution of sodium silicate gels (Garcı́a-Ruiz et al. 2003;
tubular minerals that mimic some fossilized Fig. 1b). Moreover, organic compounds produced
microbial mat textures (Pinti et al. 2009). In addi- by the abiogenic breakdown of iron carbonate can
tion, there is evidence for postdepositional coloni- condense onto these biomorphs during mild
zation of micro-cracks in the Apex Chert, and heating, thereby mimicking both the morphologi-
taken together these processes point to several cal and chemical signatures of 3.5 Ga “micro-
sources of nonindigenous carbonaceous material fossils” (Garcı́a-Ruiz et al. 2003). Perhaps more
within this unit. In summary, the Apex Chert has robust microfossil evidence may come from the
proven to be a highly controversial, but also younger Sulphur Springs Group at 3.24 Ga in the
form of pyritic filaments from a deep-sea, subseafloor is given (for a more complete review,
volcanogenic, massive sulfide deposit, interpreted see Thorseth 2011), and then these are compared to
as the fossilized remains of thermophilic, mineralized, tubular structures from the Archean.
chemotrophic prokaryotes (Rasmussen 2000; Given that pillow lavas constitute an estimated 99%
Fig. 1f). These straight, curved, or sinuous fila- of Archean greenstone belts, they represent perhaps
ments exhibit putative biological behavior includ- the largest potential habitat for seeking traces of
ing preferred orientations, clustering, and early Archean life.
intertwining. They are found in an early chert Trace of ▶ endolithic microbes has been
fabric in a subsurface drill core that is crosscut reported from both the modern oceanic crust
by later fractures. Thus, these filaments appear to and older seafloor fragments; for a comprehen-
satisfy criteria for the ▶ syngenicity and sive review, see Furnes et al. (2008). These are
▶ biogenicity of candidate microfossils and microtubular and granular cavities found at the
await supporting lines of geochemical evidence. interface of fresh and altered glass, along frac-
Siliciclastic sediments such as sandstones, tures in the rims of pillow basalts and around the
siltstones, and mudstones have also been some- margins of volcanic glass fragments. These are
what overlooked in the search for Archean traces both texturally and chemically distinct from abi-
of life. A recent study by Javaux et al. (2010) otic alteration textures found in ▶ basalts and
reports large, hollow spherical organic-walled include diverse tubular shapes such as spiraled,
microfossils known as ▶ acritarchs from the annulated, and branched forms (McLoughlin
3.2 Ga ▶ Moodies Group of South Africa. et al. 2009). Studies of recent material have
These structures pass syngenicity and found nucleic acids and bacterial and archaeal
endogenicity tests and appear to be the oldest RNA concentrated within these microborings.
and largest organic-walled spheroidal microfos- These textures may later be mineralized by
sils reported to date. They may record a plank- ▶ clays and iron oxyhydroxides that can preserve
tonic ecosystem contemporaneous with benthic localized enrichments in C, N, and P along the
▶ microbial mat textures described by Noffke margins that are interpreted as decayed cellular
et al. (2006) and will reinvigorate the search for remains. Quantitative studies of the distribution
traces of Archean life in siliciclastic sediments and abundance of these alteration textures with
with low-organic carbon contents. depth in the modern oceanic crust have found that
in the upper 350 m of the crust the granular type
Microborings is dominant. Meanwhile, the tubular alteration
Microborings are micron-sized cavities created by textures constitute only a small fraction of the
the activities of rock-dwelling microorganisms total zone of alteration and show a clear maxi-
termed endoliths. Microborings have long been mum at 120–130 m depth corresponding to
known from Precambrian silicified carbonates and lower temperatures of 70 C and thermophilic
have more recently been reported from the glassy metabolisms. Comparisons of seafloor and drill
margins of pillow lavas from modern to Archean core samples of different ages suggest that
volcanic rocks (Staudigel et al. 2008). A rock- bioalteration commences early soon after crystal-
dwelling mode of life in the Archean subseafloor lization of the basalt flows.
may have offered many attractions including prox- In the Archean tubular bioalteration, textures
imity to geothermal heat, a source of reductants, have been reported from the formerly glassy rims
principally Fe and Mn which are abundant in of pillow basalts and inter-pillow breccias from
basalts, and access to both oxidants and carbon both South Africa (Fig. 1a) and West Australia
sources carried by circulating fluids. Such habitats (Furnes et al. 2007). Some of the best examples
would also have offered protection from the ele- come from the 3.46 Ga Hooggenoeg Complex
vated UV radiation and meteoritic and cometary of South Africa and are mineralized by titanite
impacts on the early earth. First, a brief overview of (CaTiO3) that ensured preservation of the tex-
what is known about these organisms in the modern tures when the host glass was transformed to a
greenschist facies metamorphic mineral assem- Precambrian stromatolites of widely regarded

blage. These mineralized tubular structures are biogenic origin fail to qualify. The task of
1–10 mm wide, up to 200 mm long, and extend distinguishing biogenic from abiogenic stromat- A
away from “root zones” of fine-grained titanite olites is unfortunately, especially, difficult in the
that is associated with fractures in the basaltic Archean where diagenetic recrystallization and
glass (Fig. 1a). These microtubes can have a low-grade metamorphism can destroy any
segmented appearance caused by overgrowths organic micro-textures that were once present.
of metamorphic chlorite. Morphologically com- The oldest putative stromatolites are reported
parable microtubular structures have also been from the 3.49 Ga Dresser Formation of the
reported from inter-pillow breccias within the West Australia, including wrinkled planiform
3.35 Ga Euro Basalt Fm of West Australia surfaces, broad domes, and columnar forms.
(Furnes et al. 2007). These are also infilled with These occur at several localities in the North
titanite that has been dated directly using U-Pb Pole Dome both in syn-depositional barite
systematics, which confirm that the microborings mounds and hydrothermal dikes and in silicified
formed prior to an Archean 2.7 Ga phase of and hydrothermally altered ferruginous carbon-
▶ metamorphism (Banerjee et al. 2007). Late ates (Wacey 2009). They are of disputed biolog-
Archean microborings have now also been ical origin and are discussed elsewhere in this
described from 2.5 Ga pillow lavas of Wutai, volume. Some of the next oldest putative stro-
China. matolites are described from the 3.4 Ga Strelley
In summary, microborings provide an impor- Pool Chert of West Australia. The discovery of
tant tool for mapping the deep subseafloor bio- large coniform stromatolites with rare flank struc-
sphere that may represent one of the earliest tures and domal and laterally linked
habitats for life on earth and perhaps other plan- pseudocolumnar morphologies leads to a biolog-
etary surfaces. ical origin for these structures being advanced
(Hofmann et al. 1999). Subsequently, detailed
Stromatolites and Wrinkle Mats mapping of the stromatolites and investigation
▶ Stromatolites are the most abundant macrofos- of rare outcrops with good micro-textural preser-
sil in the Precambrian rock record and are a vation have found evidence for a spatiotemporal
volumetrically significant component of Precam- correlation between stromatolite morphology,
brian carbonate platforms. Stromatolites com- micro-fabric, and depositional environment
prise laminated, centimeter-to-decimeter-scale (Allwood et al. 2009). Regionally, however, the
domes, cones, columns, and planiform surfaces more typical, small, unbranched coniform stro-
that are built through a complex interplay of matolites of the Strelley Pool Chert do not show
physical, chemical, and biological processes pro- unambiguous biological characteristics or depth-
ducing an array of micro-fabrics and laminar controlled distribution and/or changes in mor-
geometries. The processes that lead to the growth phology with depth (Wacey 2010). In short, the
of stromatolites and how these can be identified Strelley Pool Chert includes a spectrum of stro-
in the ▶ fossil record are reviewed in detail else- matolitic structures, some of which are biogenic,
where in this volume. Here a nongenetic defini- but we are still a way from confidently
tion of a stromatolite is adopted because it can be distinguishing those that are undoubtedly bio-
difficult to demonstrate active biological partici- genic from those that are not microbially
pation in stromatolite growth. There have been mediated.
many attempts to develop stromatolite A morphological biosignature related to stro-
biogenicity criteria in an effort to distinguish matolites is that of wrinkle-mat textures or
laminated seafloor precipitates formed by purely microbially induced sedimentary structures
chemical processes from microbially mediated (MISS). These are formed by the interaction of
deposits. Most of these biogenicity criteria are benthic microbiota with physical sediment
so exacting, however, that the majority of dynamics, and some of the oldest come from the
3.2 Ga Moodies Group of South Africa (Noffke 1.78 and 1.68 Gyr. Moreover, these findings also
et al. 2006). These types of structures have been seemed to suggest an early accumulation of
described as orange-peel textures on bedding sur- atmospheric oxygen. Such biomarker studies,
faces with microscopic reticulated filaments of however, have long been surrounded by concerns
carbonaceous material among the sediment of contamination from nonindigenous hydrocar-
grain that have carbon isotopic signatures that bons, especially since the carbon isotope ratios of
are consistent with a biological origin. Older the extracted biomarkers were significantly
putative wrinkle-mat horizons are described enriched relative to the bulk sedimentary organic
from cherts of the Barberton (e.g., Walsh and matter. A recent nanoSIMS study has shown that
Lowe 1999). These comprise carbonaceous lam- the carbon isotope values of pyrobitumen
inae and wisps with examples of plastically (thermally altered petroleum) and ▶ kerogen
deformed carbonaceous fragments interpreted as contained within these rocks are strongly
microbial mat rip-up clasts (Fig. 1e) and pur- depleted in 13C, confirming that the indigenous
ported examples of filamentous microfossils. petroleum is 10–20 ‰ lighter than the extracted
These wrinkle-mat-type textures and morphol- hydrocarbon biomarkers (Rasmussen et al.
ogies are certainly very suggestive of microbial 2008). These findings are inconsistent with an
processing and arguably better preserved than indigenous origin for the biomarkers that, more-
anything hitherto reported from Western Austra- over, have carbon isotopic values that are atypical
lia. Recent studies documenting the facies and of late Archean organic matter. Thus, it appears
depth-dependent distribution of this carbona- that the biomarkers derived from these
ceous material using detailed petrography and 2.7 Ga-rich ▶ shales are not indigenous to the
elemental analysis have strengthened the case rock and are not robust evidence for
for biogenicity and argued for the presence of cyanobacteria and eukaryotes at 2.7 Ga
anoxygenic photosynthesizers 3.4 Gyr (Tice (Rasmussen et al. 2008). This in situ nanoSIMS
and Lowe 2004). approach to measuring carbon isotopes on differ-
ent carbon-bearing phases will be used to test the
Biomarker Compounds antiquity and endogenicity of other late Archean
In rocks that do not preserve cellular microbial and younger reports of biomarkers.
remains, ▶ biomarker compounds found in solu-
ble hydrocarbon fractions have been used as Carbon Isotopes
markers of specific biological pathways (Brocks Various microbial metabolisms have been argued
and Summons 2003). Such compounds are for on the basis of carbon isotopic ratios mea-
derived from lipids in cell membranes and repre- sured on organic matter contained within
sent an important source of information about the Archean rocks. These include anoxygenic photo-
diversity and evolution of life. For example, a synthesis from C isotopes in the range of 20 ‰
suite of lipid biomarkers extracted from 2.7 to 30 ‰ measured on kerogen (e.g., Tice and
Gyr organic-rich shales from Western Australia Lowe 2004) and methanogenesis from very low
included hopane and sterane compounds that C ratios of 56 ‰ measured on methane-bearing
were interpreted, respectively, as the membrane fluid inclusions (Ueno et al. 2006). These d13C
remnants of cyanobacteria, a group of organisms values are certainly consistent with life, but,
characterized by oxygen-producing ▶ photosyn- unfortunately, carbon isotope fractionation pat-
thesis, and of eukaryotes organisms that have a terns when taken alone are not a uniquely biolog-
membrane-bound nucleus and a complex cyto- ical signal. This is because there are alternative
skeleton (Brocks et al. 2003). These findings in nonbiological explanations for such light carbon
rocks of 2.7 Ga greatly extended the age of first isotopic values that need to be excluded, and
appearance of cyanobacteria previously esti- these are the source of much debate in Archean
mated at 2.15 Gyr old from fossil evidence rocks. For example, Fischer-Tropsch type (FTT)
and eukaryotes previously estimated at between reactions between CO and metals (Sherwood
Lollar et al. 2002) or the metamorphic reduction carbon were used to argue that sulfate-reducing
of siderite (van Zuilen et al. 2002) can generate bacteria had evolved by 3.49 Ga. More recent
carbon isotope fractionations that lie within the investigations of material from the North Pole A
“biological domain.” Thus, in the early rock have measured mass-independently fractionated
record, carbon isotopes need to be integrated sulfur isotopic anomalies (MIF) in these sulfides
with other isotope systems along with the geolog- that differs from their host barite (Philippot
ical context and any candidate morphological et al. 2007). These authors interpret this com-
traces of life. bined negative d34S and positive MIF signature
One of the most ancient claims for life comes of the sulfides as the product of microorganisms
from isotopically light carbon found in 3.8 Ga that disproportionate elemental sulfur and not
highly metamorphosed rocks from the island of sulfate-reducing bacteria. In contrast, Ueno
▶ Akilia off the west coast of Greenland. The et al. (2008) and Shen et al. (2009) found that
material analyzed was graphitic carbon found as these same microscopic sulfides possessed D33S
inclusions within grains of apatite, with a d13C values. They used these data together with D33S
signature of 20 ‰ to 50 ‰ (Mojzsis and D36S relationships to argue that their sulfides
et al. 1996). The vigorous debate that has formed dominantly by microbial sulfate reduc-
surrounded these observations provides an illus- tion. These conflicting conclusions may in part be
trative case study of the need to understand the due to methodological differences between the
complete geological history of a rock argued to studies. An alternative approach has been taken
contain chemical traces of life. Different workers by Wacey et al. (2010) who investigated mm-
have subsequently challenged the evidence for sized, diagenetic pyrite grains from a 3.4 Gyr,
life in the Akilia rocks on the basis of the age of regionally extensive shallow marine sandstone
the outcrop, the apatite petrography, and the fact unit. They reported high-resolution multiple
that the protolith is not sedimentary in origin. S isotope analysis (32S, 33S, 34S) by secondary
A parallel debate has played out regarding the ion mass spectrometry and both nanoSIMS and
original biogenic interpretation of isotopically traditional large-radius ion microprobe to reveal
light graphite in apatite crystals from another d34S values between 12 ‰ and +6‰ and D33S
site in the Isua Greenstone Belt of Greenland values between 1.65 ‰ and +1.43 ‰, from
(Mojzsis et al. 1996). Here, petrographic and pyrite grains within a single thin section.
geochemical studies have also rejected the origi- A large spread of d34S values over only 5–10
nal biogenic interpretation and proposed that the mm, together with the spatial association of pyrite
metamorphic decomposition of ferrous carbonate with C and N, indicates biological processing of
(siderite) is the more likely source of the depleted sulfur. The presence of both +D33S and -D33S
carbonaceous material (van Zuilen et al. 2002). signals overprinted by significant mass-
There does remain, however, one locality in the dependent d34S fractionation in this pyrite popu-
Isua region where the association of graphite with lation indicates that both microbial sulfate reduc-
metasedimentary rocks may still be suggestive of tion of aqueous sulfate (D 33S) and microbial
life (Rosing and Frei 2004), and this occurrence disproportionation of elemental sulfur (+D33S)
awaits further verification. were co-occurring in an open-marine,
sedimentary-hosted ecosystem in the early
Sulfur Isotopes Archean. A parallel sulfur isotope story is starting
The earliest sulfur isotope evidence suggestive of to emerge from rocks of the ▶ Barberton Green-
life comes from microscopic sulfides contained stone Belt of South Africa. Shales and black
within barite crystals in the 3.49 Gyr Dresser cherts from the 3.3 Ga Mendon Formation
Formation of North Pole, Western Australia yield d34S values with 12 ‰ variation that was
(Shen et al. 2001). Fractionations of up to argued to be greater than that expected from
21.1 ‰ between the sulfides and coexisting sul- purely magmatic or hydrothermal H2S and due
fates together with the co-occurrence of organic to bacterial sulfate reduction (Ohmoto
et al. 1993). But as yet, no corresponding MIF and carbonates and also fine-grained siliciclastics
signature suggestive of sulfur-disproportionating to locate organic-walled microfossils and stro-
bacteria has been reported from the Barberton matolitic structures in a range of lithologies in
rocks. addition to classical carbonates. In conclusion,
In late Archean rocks, sulfur isotope evidence our current understanding of Archean ecosystems
in conjunction with carbon isotopes and rare earth is like an unfinished and jumbled-up jigsaw – as
element studies give more definitive evidence for new techniques, preservational windows, and
the emergence of sulfur metabolisms. Investiga- rock types are found, new pieces in this jigsaw
tions of the 2.7–2.6 Ga Belingwe Greenstone Belt of early life and environments will come together
of Zimbabwe have found pyrites in sulfidic shales refining our global picture.
with a wide range of d34S values from 21.1 ‰
to +16.7 ‰. This range together with the pyrite
morphology and isotopic heterogeneity provides See Also
good evidence for sulfate-reducing and possible
sulfur-oxidizing bacteria at this time (Grassineau ▶ Abiotic
et al. 2001). Carbon isotopic investigation ▶ Akilia
accompanied by rare earth element analysis of ▶ Apex Chert
associated stromatolitic and non-stromatolitic ▶ Archean Drilling Projects
sediments across an onshore-offshore gradient ▶ Archean Environmental Conditions
has also been used to argue for the presence of a ▶ Archean Eon
diverse microbial ecosystem including ▶ Archean Tectonics
anoxygenic photosynthesizer, ▶ methanogens, ▶ Barberton Greenstone Belt
and ▶ methanotrophs at this time (Grassineau ▶ Biogenicity
et al. 2001). This type of integrated approach is ▶ Biomarkers
the best way of deciphering Archean traces ▶ Biomarkers, Morphological
of life. ▶ Carbon Isotopes as a Geochemical Tracer
▶ Chemolithoautotroph
▶ Complexity
Future Directions ▶ Cyanobacteria
▶ Earth’s Atmosphere, History of the Origins
The controversies that currently surround the ear- ▶ Endogenicity
liest claims for life on earth help astrobiologists ▶ Endolithic
to develop criteria for testing new and existing ▶ Fischer-Tropsch Effects on Isotopic
claims for extraterrestrial life. Some new tech- Fractionation
niques and approaches to seeking and verifying ▶ GC/MS
Archean traces of life are now highlighted. ▶ Geochronology
Firstly, emerging nanoscale techniques that ▶ Greenstone Belts
allow high-resolution elemental and isotopic ▶ Isotope Biosignatures
analysis using nanoSIMS and synchrotron-based ▶ Isotopic Ratio
techniques give the opportunity to investigate ▶ Isua Supracrustal Belt
putative microbial fabrics at a scale never previ- ▶ Kerogen
ously obtainable and will help assess the ▶ Metasediments
biogenicity of stromatolites and microfossils in ▶ Microbial Mats
particular (e.g., Wacey 2009 and references ▶ Microfossils
therein). Secondly, renewed interest is being ▶ Nitrogen Isotopes
paid to a wider range of rock types in the search ▶ North Pole Dome (Pilbara, Western Australia)
for Archean traces of life including microborings ▶ Oxygen Isotopes
in not just volcanic glass but also silicate minerals ▶ Photosynthesis
▶ Pilbara Craton variations on the origins of early Archean BIFs from

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Areology (Greek, Ares “Mars” and logos
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Y (2006) Evidence from fluid inclusions for microbial ▶ Mars, excluding its atmosphere. It comprises
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properties, dynamics, origin, and evolution of
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mation: new evidence for microbial sulfate reduction shaped Mars. The term areology is not frequently
in the early Archean. Geochim Cosmochim Acta used because scientifically and methodologically
72:5675–5691
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it is synonymous to the term “Geology of Mars.”
(2007) Earth’s oldest rocks: developments in Precam- Thus it is common to use the latter term instead of
brian geology, vol 15. Elsevier, London “areology.”
Argonium 155
See Also 174.21. Its three-letter symbol is Arg and

one-letter symbol is R. It has a guanidinium
▶ Chronostratigraphy group (H2NC(=NH)NH2) in its side chain. The A
▶ Geological Time Scale, History of side chain is basic with a pKa of 12.48. When
▶ Mars protonated, the positive charge of the
▶ Mars Stratigraphy guanidinium ion is delocalized on the three nitro-
▶ Selenology gen atoms. The isoelectric point (pI) of
arginine is 10.76, which is the highest among
the protein amino acids. Adult humans are able
to biosynthesize arginine, but infants cannot.
Thus, it is an essential amino acid only for
Argentina Space Agency
infants. To date, arginine has not been found in
extraterrestrial bodies like carbonaceous
▶ CONAE, Argentina
chondrites.
Argillaceous Earth See Also

▶ Clay
▶ Amino Acid
▶ Protein
Arginine
Kensei Kobayashi Argonium

Hodogaya-ku, Yokohama, Japan William M. Irvine
Definition
Synonyms
Arginine, shown in Fig. 1, is one of the 20 ▶ pro-
tein ▶ amino acids. Its molecular weight is
ArH+
Arginine, NH
Fig. 1 Structural formula H2N Definition
of arginine
NH Argonium is the argon hydride cation, ArH+.
Argon has three naturally occurring isotopes,
with the cosmically most abundant being 36Ar,
followed by 38Ar, and then 40Ar. On Earth, in
H contrast, 40Ar is by far the most abundant isotope,
O derived primarily from the radioactive decay of
40
H2N K. The K-Ar decay is important for geochro-
nology. Argonium is the first noble gas molecule
OH to have been found in space, with both the 36Ar
156 ArH+
and the 38Ar isotopologs observed by astrono-

mers. Extensive laboratory studies of the rota- ArH+
tional spectra and the electric dipole moment
have been carried out for both 36ArH+ and ▶ Argonium
38
AH+; AH+ is quite stable, with a dissociation
energy of almost 4 eV.
History
Ariel
Astronomically, argonium was first reported in
the Crab Nebula, the remnant of the supernova Therese Encrenaz
observed by Chinese astronomers/astrologers in LESIA, Observatoire de Paris - Section de
1,054 CE (Barlow et al. 2013). The observed 36Ar Meudon, Meudon, France
was presumably produced by ▶ explosive nucle-
osynthesis during the supernova event. Both
36
ArH+ and 38AH+ were more recently reported
toward the Galactic Center molecular cloud Definition
SgrB2 (Schilke et al. 2014). Because ArH+ reacts
rapidly with molecular hydrogen, H2, but not at Ariel is one of the five big satellites of ▶ Uranus
cold interstellar temperatures with atomic and the closest to the planet. It was discovered by
hydrogen, H, Schilke et al. suggest that ArH+ William Lassen in 1851. Its diameter is 1,160 km
may well be a useful tracer of the neutral atomic and its distance to Uranus is 191,000 km or 7.5
hydrogen in galaxies. planetary radii. Its density is 1.66 g/cm3.
Ariel has been explored by the Voyager
2 spacecraft which flew by the Uranian system
in January 1986. Ariel is assumed to consist
See Also of about 30 % silicates and 70 % ices. It has a
bright surface that shows a network of
▶ Geochronology canyons and faults, such that, after
▶ Isotopolog Miranda, Ariel is the most geologically active
▶ Molecular Cloud among Uranus’ satellites. The longest can-
yon (622 km) is Kachina Chasmata. The activity
References and Further Reading may result from tidal heating due to the
proximity of Uranus at the time of the satellite’s
Barlow MJ, Swinyard BM, Owen PJ, Cernicharo J, formation.
Gomez HL, Ivison RJ, Krause O, Lim TL,
Matsuura M, Miller S, Olofsson G, Polehampton ET
(2013) Detection of a noble gas molecular ion, 36ArH+,
in the crab nebula. Science 342:1343–1345
Schilke P, Neufeld DA, M€ uller HSP, Comito C, Bergin
EA, Lis DC, Gerin M, Black JH, Wolfire M, See Also
Indriolo N, Pearson JC, Menten KM, Winkel B,
Sánchez-Monge Á, Möller T, Godard B, Falgarone ▶ Giant Planets
E (2014) Ubiquitous argonium (ArH+) in the diffuse
interstellar medium – a molecular tracer of almost pure ▶ Uranus
atomic gas. Astron Astrophys 566:A29 ▶ Voyager, Spacecraft
Arrhenius Svante 157
function of temperature could be described by the

Aromatic Hydrocarbon equation
A
Henderson James (Jim) Cleaves II ln k ¼ ln A Ea =RT
Institute of Technology, Meguro–ku, Tokyo, where k is the reaction rate, R is the universal gas
Japan constant, Ea is the ▶ activation energy (the energy
Institute for Advanced Study, Princeton, NJ, required in order for the reactants to react), T is the
USA absolute temperature (in degrees K), and A is the
Blue Marble Space Institute of Science, so-called pre-exponential factor (associated with
Washington, DC, USA collision and transition state theory).
Center for Chemical Evolution, Georgia Institute A plot of ln k versus 1/T often yields a straight
of Technology, Atlanta, GA, USA line, the slope of which is equal to the activation
energy of the reaction divided by the universal
Definition gas constant (Ea/R) and the y-intercept of which
is equal to ln A.
An aromatic hydrocarbon is a cyclic hydrocarbon
where the series of saturated and unsaturated carbon-
See Also
carbon bonds satisfies H€uckel’s rules, i.e., where the
number of electrons in double and triple bonds in the
▶ Activation Energy
ring is 4n + 2, where n = 0 or any positive integer.
The name derives from the fact that the first such
molecules discovered tended to have an aromatic
odor. They are typically somewhat more stable than
their hydrogen-saturated analogues. Aromatic Arrhenius Svante
hydrocarbons may be monocyclic or polycyclic.
Gerda Horneck
See Also Aerospace Medicine, Radiation Biology,
Köln, Germany
▶ Benzene
▶ PAH
History
Svante August Arrhenius (1859–1927), Swedish

Arrhenius Plot scientist, received the Nobel Prize in Chemistry
in 1903 “in recognition of the extraordinary ser-
Jeffrey Bada vices he has rendered to the advancement of
Scripps Institution of Oceanography, La Jolla, chemistry by his electrolytic theory of dissocia-
CA, USA tion.” Among other achievements, Arrhenius is
famous for the Arrhenius equation, which gives
Definition the dependence of the rate constant k of a chem-
ical reaction on the temperature T (in K) and the
In 1889, the Swedish chemist Svante Arrhenius activation energy of the reaction. For astrobiolo-
showed that the rate of a chemical reaction as a gists, Arrhenius is famous for his thoughts that
158 Artificial Cells
microscopic forms of life, for example, ▶ spores, Synonyms

can be propagated in space, driven by the radia-
tion pressure from the Sun and thereby seeding Reaction network
life from one planet to another or even between
planets of different stellar systems. Arrhenius
based his considerations on the fact that the
space between the planets of our Solar System is Definition
teeming with micron-sized cosmic dust particles,
which at a critical size below 1.5 mm would be Artificial chemistry (AC) is a field of research
blown away from the Sun with high speed pushed that studies systems that are similar to, and
by radiation pressure of the Sun. Herewith Arrhe- commonly a generalization of, chemical net-
nius provided a scientific rationale for the theory works. Those studies are usually done through
of ▶ Panspermia, now called Radiopanspermia. computer simulations which are also called arti-
ficial chemistries. An AC can be defined as triplet
{M, R, A}, with M being the set of possible
See Also molecules (sometimes infinite), R the set of pos-
sible reactions, and A the algorithm (Dittrich
▶ Arrhenius Plot et al. 2001).
▶ Lithopanspermia
▶ Panspermia
▶ Spore
Overview
References and Further Reading Artificial chemistry grew as a field from the early
Arrhenius S (1903) Die Verbreitung des Lebens im
1990s. Different aims converged in producing
Weltenraum. Umschau 7:481–485 this field.
Arrhenius S (1908) Worlds in the making: the evolution of One of the aims of artificial chemistry was to
the universe. Harper & Row, New York investigate evolution and the appearance of life.
Some of the main questions that have been inves-
tigated deal with biology, proto-biology, and evo-
Artificial Cells lution and take a bottom-up approach to
▶ artificial life:
▶ Cell Models
▶ Protocell • Can we generate an artificial chemistry that
can generate an artificial life?
• What kind of chemistry can sustain life?
• How does the Darwinian evolutionary process
Artificial Chemistries depend upon the chemistry on which it is
based?
Pietro Speroni di Fenizio
CISUC, Department of Informatics Engineering, Of course, as we extrapolate out of chemistry
University of Coimbra, Coimbra, Portugal the basic network of relations that can produce
life, this would eventually help to recognize life
in different contexts.
Keywords Another, more general, aim was to study
reaction networks. Reaction networks appear in
Artificial chemistries; Artificial life; Evolution; multiple fields, but our ability to study them
Reaction network has generally been limited by a difficulty in
Artificial Chemistries 159
solving differential equations where the Basic Methodology

number of different interacting elements
(diversity, i.e., number of equations) were The methodology is slightly different for Spatial A
too high, and the quantity of each participating Artificial Chemistries and for Complex Artificial
element in the reaction network (population size) Chemistries.
was too low (which could lead to an Complex Artificial Chemistries, as mentioned
element disappearing and the set of reactions before, are essentially reaction networks with a
changing). Some artificial chemistries high diversity and a low population size. Such an
investigated exactly this problem. As such the AC can be defined as a triplet {molecules, reac-
aim of artificial chemistries has often been a tions, algorithm}. Reactions are usually binary
qualitative study, looking for what sets of mole- (i.e., two molecules reacting and generating a
cules would be present in the reactor, more than third). Yet not all pair of molecules can react.
how many elements would be present for each When two molecules cannot react, they are said
type of element in the system (Fontana and to be elastic. The basic algorithm is generally
Buss 1996). very simple:
Due to the computational difficulty in simu-
lating a chemical system that is both unbounded 1. Define a Soup S as a multiset of m molecules
in the complexity of the molecules involved and out of M.
that permits a faithful representation of space, the 2. Two molecules are randomly selected.
field tends to be divided into two main subfields: 3. If the two molecules can react, a reaction takes
Complex Artificial Chemistries and Spatial Arti- place:
ficial Chemistries. 1. The result of the reaction is inserted in
Complex Artificial Chemistries are the the Soup.
most simple type of system and assume a 2. A random molecule is extracted from the
well-mixed reactor. Each molecule is either Soup and eliminated.
present or absent, with no information about 4. Go to 2.
the position. Pairs of molecules are then ran-
domly selected and reacted. Such a simple In many Artificial Chemistries, the reactions
algorithm frees the computational resources are catalytic, so when two molecules react, they
permitting the presence of complex are not taken away from the Soup while their
molecules, with different behaviors. It is not product is added to the Soup (3a). Instead, the
uncommon for artificial chemistries to study molecules catalytically induce the formation of
the behavior of little program strings with com- the new molecule out of a substrate of basic
putational capabilities (traditionally lambda material (too vast and ubiquitous to be explicitly
terms). modeled), while the disappearance (3b) would
Spatial Artificial Chemistries traditionally model the outflux due to molecules being washed
study systems made up of many simple elements, away or breaking apart (Dittrich et al. 2001). This
floating in a 2D or in a 3D spatial environment. traditional structure has been strongly criticized
The basic elements (in some systems called (e.g., for not considering conservation of mass),
atoms) can often link together creating long and many alternatives have been offered but gen-
chains (usually called molecules). In the first erally with only partial differences in the
pioneering works of Spatial Artificial Chemis- observed behavior. Traditionally the standard
tries, the positions of the molecules were impre- way to investigate such a system is to run it
cise, as each molecule was generally assigned to a until no new molecule would be generated, at
position on a lattice. Recent computational pro- which point the resulting multiset is studied.
gress permitted a more faithful representation of Lately, a more advanced method has been to
the movement using differential equations calculate all the possible sets where the system
(Fellermann 2009). could stop (called organizations) and study the
160 Artificial Chemistries
structure of the lattice of organizations and use Applications

this to map the movement of the artificial chem-
istry as time progresses (Kaleta 2009). While the long-term aim of artificial chemistry, to
Spatial Artificial Chemistries start in general investigate the appearance of life, has not been
with a set of atoms, to which a position in space reached, a number of partial findings were recog-
is randomly assigned. At each time step all the nized as being useful in different fields. As Com-
atoms are randomly moved, and when two atoms plex Artificial Chemistry can investigate reaction
end up near each other, they have the possibility of network with a high diversity, they are often the
colliding. Unless the system is superimposed on a right tool to study ▶ biological networks. In par-
lattice, the movement of the molecules would fol- ticular, studies have been done on gene regula-
low a Brownian motion style of movement or a tory networks and the internal metabolism of a
dissipative particle dynamic style, where the cell. In this regard, AC has started merging with
forces interacting on a molecule are taken into the other tools inside systems biology and bioin-
account in greater detail (Fellermann 2009). formatics. Other studies have tried to consider
social systems, language, and economical sys-
tems (Dittrich et al. 2001).
Key Research Findings
The first result in AC is that there are sets of Future Directions

molecules, called organizations, which are qual-
itatively stable, in the sense that each molecule Spatial Artificial Chemistries and Complex Arti-
present in the set can be generated by the reac- ficial Chemistries are going in separate ways,
tions inside the set and that all reactions inside the mostly answering different questions about
cell can only generate molecules that are already nature. Spatial Artificial Chemistries try to pro-
inside the cell. Such organizations form an alge- duce a minimal cell, but have not yet succeeded
braic lattice, and in the absence of external inputs, in generating a spontaneous emergence of a full
each experiment eventually leads to the system Darwinian evolutionary system. So we can
reaching one of those sets. As artificial chemis- expect more research to go in this direction.
tries can be represented using ordinary differen- Results in Complex Artificial Chemistries have
tial equations (ODE), it has been shown that fixed been more connected with systems biology, and
points in the ODE of the system exist only inside artificial chemistries have been used to explicitly
organizations. In other words, if we take a fixed study system biology models, and recent studies
point of the system, find the molecules that are show that it is possible to use AC to build pro-
present with a quantity higher than 0, and then grammable systems made up of many interacting
this set of molecules forms an organization. components. In the future, we can expect those
Those results permit a preliminary study of an two trends to merge, as scientists investigate
artificial chemistry by finding (through algebraic chemistries that can evolve and that can be
studies) the lattice of organizations and produce a programmed to evolve.
map of the system on which it is possible to track
the changes in the system (Dittrich and Speroni di
Fenizio 2007). See also
Spatial Artificial Chemistries have been suc-
cessfully used to model the lipid bilayer of cell ▶ Artificial Life
membranes and to investigate the spontaneous ▶ Autopoiesis
emergence of artificial life protocells. More ▶ Biological Networks
recently, experiments have been carried out gen- ▶ Chemical Reaction Network
erating protocells that are able to reproduce ▶ Darwin’s Conception of the Origins of Life
(Rasmussen et al. 2007). ▶ Evolution, Biological
Artificial Life 161
References and Further Reading elucidate the fundamental mechanisms common

to living organisms. It focuses on the rule-based
Dittrich P, Speroni di Fenizio P (2007) Chemical organi- mechanisms making life possible, supposedly
sation theory. Bull Math Biol 69(4):1199–1231 A
neutral with respect to their underlying material
Dittrich P et al (2001) Artificial chemistries – a review.
Artif Life 7:225–275 embodiment, and to replicate them in a
Fellermann H (2009) Spatially resolved artificial chemis- non-biochemical substrate. In artificial life, the
try. In: Adamatzky A, Komosinski M (eds) Artificial importance of the substrate is purposefully under-
life models in software, 2nd edn. Springer, p 343
stated for the benefit of the function. Minimal life
Fontana W (1992) Algorithmic chemistry. In: Proceedings
of artificial life II conference 5:159–210 begins at the intersection of a series of processes
Fontana W, Buss LW (1996) The barrier of objects: from that need to be isolated, differentiated, and dupli-
dynamical systems to bounded organizations. In: cated as such in computers. Only software devel-
Karlqvist A, Casti J (eds) Boundaries and barriers.
opment and running make it possible to
Addison-Wesley, Redwood City, pp 56–116
Kaleta C (2009) From artificial chemistries to systems biol- understand the way these processes are intimately
ogy. In: Adamatzky A, Komosinski M (eds) Artificial life interconnected in order for life to appear at the
models in software, vol 2. Springer, London, pp 319–342 crossroads.
Rasmussen S et al (2007) Life cycle of a minimal
protocell – a dissipative particle dynamics study.
Artif Life 13(4):319–345
Overview
Artificial life obviously relates to astrobiology;

Artificial Evolution
this other recent interdisciplinary field of scien-
tific research equally centered on life and the
▶ Evolution, In Vitro
study of its origins, not only on the obvious envi-
ronment of Earth but also throughout the uni-
verse. Astrobiology cannot restrict itself to a
Artificial Life mere materialistic view of life, in order to detect
it elsewhere, as the material substrate could be
Hugues Bersini something totally different. This substrate could
IRIDIA, Université Libre de Bruxelles, Brussels, be as much singular on a distant planet as it could
Belgium be in the RAM memory somewhere in a univer-
sity computer laboratory. The presence of life
might be suspected through its functions, much
Keywords before scientists are able to dissect it. Artificial
life does not attempt to provide an extra thou-
Bioinformatics; Genetic algorithms; Networks; sandth attempt at the definition of life, any more
Self-replication; Software simulations; Von than do most biologists. As a matter of fact, the
Neumann concept of “life,” as opposed to “gravity” or
“electromagnetism” or “quantum reduction of a
wave packet,” has already been in widespread
Synonyms existence prior to any scientific reading or reifi-
cation. The rejection of an authoritative definition
Life, artificial; Theoretical biology of “life” is often compensated for by a list of
functional properties that never finds unanimity
among its authors. Some demand more proper-
Definition ties, and others require fewer of those properties
that are often expressed in terms of a vague
Artificial ▶ life uses software simulation and, to a expression such as “self-maintenance,” “self-
lesser degree, robotics in order to abstract and organization,” “metabolism,” “autonomy,”
162 Artificial Life
“▶ self-replication,” and “open-ended evolu- them, the more the predictions will move from
tion.” A first determining role of artificial life qualitative to precise and the easier the model
consists in the writing and implementing of soft- will be to validate according to the Karl Popper
ware versions of these properties and of the way ideal falsifying process.
they do connect, so as to render them unambigu- Finally, through systematic software experi-
ous, making them algorithmically precise enough ments, these platforms can lead to the discovery
that, at the end, the only reason for disagreement of new natural laws, whose impact will be greater
on the definition of life would lie in the length or if the simulated abstractions will be present in
the composition of this list and on none of its many biological realms. In the 1950s, when
items. Alan Turing (1952) discovered that a simple dif-
The biologist obviously remains the most fusion phenomenon propagating itself at different
important partner; but what may he expect from speeds, depending on whether it is subject to a
this “artificial life”? These computer platforms negative or positive influence, produces zebra or
could be useful in several ways, presented in the alternating motifs, it had a considerable effect on
following in terms of their increasing importance a whole section of biology studying the genesis of
or by force of impact. First of all, they can open forms (animal skins, shells of sea creatures
the door to a new style of teaching and advocating (Meinhardt 1998)). When some scientists discov-
of the major biological ideas, that is, computer ered that the number of attractors in a Boolean
software as pedagogical help, as, for example, network or a neural network exhibits a linear
Richard Dawkins (1986) who, bearing the Dar- dependency on the number of units in these net-
winian good news, did so with the help of a works (Kauffman 1993, 1995), these results are
computer simulation where sophisticated crea- equally well applied to the number of cells
tures known as “biomorphs” evolve on a com- expressed as dynamic attractors in a genetic net-
puter screen by means of a genetic algorithm. work or the quantity of information capable of
These same platforms and simulations can, inso- being memorized in a neural network. Entire
far as they are sufficiently flexible, quantifiable, chapters of biology dedicated to networks
and universal, be used more precisely by the (neural, genetic, protein, immune, hormonal)
biologist, who will find in them a simplified had to be rewritten in the light of these discover-
means of simulating and validating a given bio- ies. When some scientists recently observed a
logical system under study. Cellular automata, nonuniform connectivity in many networks,
Boolean networks, ▶ genetic algorithms, and whether social, technological, or biological,
algorithmic chemistry are excellent examples of showing a small number of key nodes with a
software to download, parameterize, and use to large number of connections and a greater num-
produce the natural phenomena required. Their ber of nodes with far fewer, and when, in addi-
predictive power varies from very qualitative tion, they explained the way in which these
(their results apparently reproduce very general networks are built in time (Barabasi 2002) by
trends of the real world) to very quantitative (the preferential attachment, again biology was
numbers produced by the computer may be pre- clearly affected. Artificial life is of course at its
cise enough to be compared with those measured apogee when it reveals new biological facts,
in the real world). Although being at first very destabilizing the presuppositions of biologists or
qualitative, a precise and clear coding is already generating new knowledge, rather than simply
the guarantee of an advanced understanding illustrating or refining the old.
accepted by all. Algorithmic writing is an essen- In the next section, I shall attempt to set out the
tial stage in formalizing the elements of the history of life as the disciples of artificial life
model and making them objective. The more the understand it, by placing the different landmark
model allows us to integrate what we know about steps on a temporal and causal axis, showing
the reality being reproduced, that is, the detailed which one is indispensable to the appearance
structures of objects and relationships between of the next and how it connects to the next.
Artificial Life 163
This history will certainly be very incomplete and compensating and reestablishing any disruption
full of numerous unknowns, but most people in concentration undergone by anyone of them.
involved in artificial life will be in agreement. The bigger the network, the more stable it should A
They will mainly disagree on the number of be and the more molecules it will maintain in a
these functions and on the causal sequence of concentration zone that will vary very little,
their appearance, acknowledging, however, that despite external disruptions.
the appearance of any would have been condi- A network of this kind will be materially closed
tioned by the presence and the functioning of the but energetically open if none of the molecules
previous ones. The task of artificial life is to set appears in or disappears from the network as a
up experimental software platforms where these result of material fluxes, whereas energy, originat-
different lessons, whether taken in isolation or ing in external sources, is necessary for the reac-
together, are tested, simulated, and, more system- tions to start and take place. The presence of such
atically, analyzed. I shall sketch some of these an energy flux, maintaining the network far from
existing software platforms whose running the thermodynamic equilibrium, is needed, since,
delivers interesting take-home messages to without it, no reactive flow would be possible
open-minded biologists. circulating through the entire network.
A molecular end of the cycle must be reenergized
The History of Life as Seen by Artificial Life in order to start again the whole circular reaction
Proponents process. This cycle thus acts as a chemical
machine, energetically driven from the outside.
Appearance of Chemical Reaction Cycles and As soon as one of the molecules is being produced
Autocatalytic Networks in the network without, in its turn, producing one
In order for a system to emerge and maintain of the molecules making up the network, it absorbs
itself inside a soup of molecules that are poten- and thus destroys the network. In the presence of
tially reactive and contain very varied constitu- molecules of this kind, produced but nonproduc-
ents (which could correspond to the initial tive (a kind of waste), the only way of maintaining
conditions required for life to appear, i.e., in the the network becomes to feed it materially and to
primordial soup), this reactive system must form make it open to material influx. The network acts
an internally cycled network or a closed organi- on the flow of material and energy as an interme-
zation, in which every molecule is consumed and diate ongoing stabilization zone, made up of mol-
reproduced by the network. Above all, in order ecules that may be useful to other vital functions
for life to begin, all of the constituent components (such as the composition of enclosing membranes
must have been able to stabilize themselves in or catalyzing self-replication), to be described in
time. These closed networks of chemical reac- the following sections. It transforms, as much as it
tions are thus perfect examples of systems, “keeps on,” all the chemical agents that it recruits.
which, although heterogeneous, are capable of Biologists generally agree that a reactive network
maintaining themselves indefinitely, despite the must exist prior to the appearance of life, at least to
shocks and impacts that attempt to destabilize catalyze and make possible the other life processes
them. This comes about through a subtle self- such as genetic reading and coding; it is open to
regeneration mechanism, where the molecules external influences in terms of matter and energy
end up producing those molecules that have pro- but necessarily contains a series of active cycles.
duced them. It may be obtained on a basic level in They are most often designated as “▶ metabo-
a perfectly reversible chemical reaction but can lism” or “proto-metabolism,” the most popular
be obtained more subtly in the presence of a lot of and active advocates of this “metabolism-first”
intermediary molecules and catalysts. By this hypothetical scenario of the origin of life being
reaction-based roundabout in which they all par- (De Duve 2002; Ganti 2003; Maynard Smith and
ticipate, all molecules contribute to maintaining Szathmary 1999; Kauffman 1993; Shapiro 2007;
themselves at a constant concentration, Dyson 1999).
164 Artificial Life
Artificial Life,
Fig. 1 Representation of a
network of chemical
reactions of polymerization baaaa
(a + b ! ab) and baaaaaab
depolymerization
(ab ! a + b) taking place in
a simulated chemical
reactor. Molecules are aab
baa
represented by circles and
reactions by square. Each aa
reaction can be catalyzed,
like the arrows pointing to aaab
the squares show, by a
molecule of the network a
(giving rise to an ba baaa
autocatalytic network).
Some molecules can appear
(like the molecule “aab”) or
simply disappear from the
baa
network. Reaction cycles b
can appear, like the one
surrounded in the figure
(aa ! baaaa ! baaaaaab
! baaa ! aa) bb
babb
aab
baaa
babbbb
Lenaerts and Bersini (2009) give priority to concentration vanishes in time. Both the structure
the study of chemical reaction networks, viewing of the network and the concentration of its con-
them as key protagonists in the appearance of life. stituents tend to stabilize over time. Kauffman
These chemical reaction networks, where the (1993, 1995) and Fontana (1992) were the fore-
nodes are the molecules participating in the reac- runners in the study of the genesis and properties
tions and the connections are the reactions of these networks. Figure 1 illustrates the work of
linking the reacting molecules to the molecules these two artificial life pioneers, dedicated to the
produced, are generally characterized by fixed study of prebiotic chemistry, limiting the reac-
point dynamics, the chemical balances during tions studied to polymerization, such as aa + bb
which the producers and the products mutually ! aabb, or, inversely, depolymerization or
support each other. The attractors in which these hydrolysis, such as abaa ! ab + aa.
networks fix themselves are as dynamic (the con- Kauffman showed that provided the probabil-
centrations slowly stabilize) as they are structural ity that a reaction takes place is affected by the
(the molecules participating in the network are presence of a catalyst, which is itself produced by
chosen and “trapped” by the network as a whole). the network (in such a case the whole network is
These networks are perfect examples of systems said to be autocatalytic), a phenomenon of perco-
that combine dynamics (the chemical kinetics in lation or phase transition, characteristic of this
this case) and metadynamics (the network topo- type of simulation, is produced. For probabilities
logical change), as new molecules may appear as that are too low, the network does not pop up
the results of reactions while some of the mole- because the reactions are too improbable, but as
cules in the network may disappear if their soon as a threshold value is reached for this same
Artificial Life 165
1)
2) a
b
a
b
3)
a b
OR
Artificial Life, Fig. 2 The OO chemical simulator devel- right, the outcome of the simulator is an evolving reaction
oped by Lenaerts and Bersini (2009). On the left, the network, which can be studied in its own right (the pres-
molecules are represented as canonical graphs. On the ence of cycles, the type of topology)
probability, the network “percolates,” giving rise are also capable of self-regeneration and self-
to multiple molecules produced by multiple reac- replication.
tions. Kauffman grants a privileged status to this Lenaerts and Bersini (2009) have programmed
threshold value and to the giant “explosive” net- the genesis of these chemical reaction networks
work resulting from it (in his scenario of the by adopting the object-oriented (OO) program-
origin of life), without really arguing the reason ming paradigm. The OO simulator aims to repro-
why such a status should exist, but passing the duce a chemical reactor and the reaction network
immense interest and enthusiasm that the phe- that emerges from it (like that shown in Fig. 2).
nomena of phase transitions arouse among phys- This coevolutionary (dynamics + metadynamics)
icists onto the world of biology. Fontana for his model incorporates the logical structure of con-
part is concerned with the inevitable appearance stitutional chemistry and its kinetics on the one
of reaction cycles (such as that illustrated in hand and the topological evolution of the chem-
Fig. 1). All the molecules produced by these ical reaction network on the other hand. The
cycles in the network in turn produce molecules network topology influences the kinetics and the
of the network. He is among those many biolo- other way round, since only molecules with a
gists who see these closed networks or organiza- sufficient concentration are allowed to participate
tions as forming a key stage in the appearance of in new reactions (to avoid a combinatorial explo-
life, due both to their stability and to the fact that sion of molecules and reactions). The model is
they form structural and dynamic attractors for expressed in a syntax that remains as close as
the system. They cause a stabilization and inter- possible to real chemistry. Starting with some
nal regulation zone together with an energetic initial molecular objects and some initial reaction
motor in a chemical soup, which is continually objects, the simulator follows the appearance of
being crossed by a flow of matter and energy. new molecules and the reactions in which they
Fontana goes on to show how these networks participate, as well as the development of their
166 Artificial Life
concentration over a period of time. The mole- Chemical reactions

cules are coded as canonical graphs. They are a h
made up of atoms and bonds that open, close, or L L∗ LL L+ L
a βh
break during the reactions. The result of the sim- D D∗ DL L+ D
ulation consists in various reaction networks,
b βh
unfolding in time and whose properties can be L∗ L LD L+ D
further studied (for instance, the presence and the b h
D∗ D DD D+ D
properties of reaction cycles or the nature of the
p e
network-particular topology such as scale-free or L∗ + L LL LD DD
random). ap γe
One of these reaction schemes, in addition to D∗ + L DL DD LD
just cycling, can also be ▶ autocatalytic, when a ap γe
L∗ + D LD LL DL
product of the reaction cycle has twice the con-
p e
centration of one of the reactant: a + b ! a + a. D∗ + D DD DL LL
This is, for instance, the case of the so-called
formose reaction (that Ganti and Szatmary have Reaction network
discussed at large in Ganti 2003), during which a e e
two-carbon molecule, reacting twice with LL DL LD DD
a monomer composed of one carbon, leads to a γe βh γe
four-carbon molecule, which then splits in order αp αp
h h
to duplicate the original molecule. This is the
chemical variant of genetic self-replication,
since in both cases an original molecule is dupli- p L D p
cated. As will be discussed later, Ganti has been
the first to connect and synchronize these two b a a b
replication processes, chemical and genetic, in
order for the cell to simultaneously duplicate its
boundary, its metabolism, and its informational L∗ D∗
support. In the presence of autocatalysis, the reac-
Artificial Life, Fig. 3 The prebiotic chemical reactor
tion kinetics amounts to an exponential increase system responsible for a homochiral steady state studied
and, more interestingly, when various autocata- by Plasson et al. (2007). The complete set of reactions is
lytic cycles enter in antagonistic interaction, turns indicated, containing activation (the necessary energy
source), polymerization and hydrolysis (which together
out to be responsible for symmetry breaking (one
shape the cycles), and epimerization (which induces the
of the cycle, initially favored, wins and takes it competition between the enantiomers)
all). The early origin of life should not be studied
without taking account of the self-organization of
chemical networks, the emergence and antago-
nism of autocatalytic cycles, and how energy same type of polymerization and depolymeriza-
flows drive the whole process. Such chemical tion reactions as the one studied by Fontana. In
networks are, for instance, appealing to the effort the additional presence of epimerization reac-
to understand the onset of biological tions allowing the transformation of a right-
homochirality as the destabilization of the race- hand monomer into a left-hand one and vice
mic state resulting from the competition between versa, the concentration of one family of mono-
enantiomers and from amplification processes mers (for instance, the left one) vanishes in favor
concerning both autocatalytic competitors (one of the other. The flux of energy is transferred and
left oriented and the other right; see Plasson efficiently distributed through the system, lead-
et al. 2007). The chemical reaction network ing to cycle competitions and to the stabilization
under study (shown in Fig. 3) is made up of the of asymmetric states.
Artificial Life 167
Production by This Network of a Membrane components trapped during its formation. They
Promoting Individualization and Catalyzing do, however, actively channel in and out the most
Constitutive Reactions appropriate chemicals for maintaining A
The appearance of a reaction network of this kind themselves.
undeniably creates the stability necessary for In assimilating living organisms to autopoietic
exploiting its constituents in many reactive sys- systems, Varela et al. (1974) were the first to
tems such as the ones dedicated to the construc- insist that this membrane should be endogenously
tion of ▶ membranes or the replication of produced by the elements and the reactions mak-
molecules carrying the ▶ genetic code. This net- ing up the network (e.g., lipids would come from
work also acts as a primary filter as it can accept the reactions of the network itself) and would in
new molecules within it but can equally well return promote the emergence and self-
reject other molecules seeking to be incorporated maintenance of the network. The membrane can
within it. They will be rejected, as they do not help with the appearance of the reactive and
participate in any of the reactions making up the growing network by the frontiers that it sets up,
network. Can we see a primary form of individ- by the concentration of certain molecules trapped
ualization in this network? No, because by defi- in it, or by acting as a catalyst to some of the
nition, it can only be unique as no spatial frontier reactions due to its geometry or its makeup. Basi-
allows it to be distinguished from another net- cally, autopoiesis requires a cogeneration of the
work. Although it is roughly possible to conceive membrane and of the reactive network that it
of an interpenetration of several chemical net- “walls up.” The network presents a double
works, establishing a clear separation between closure – one chemical, linked to the cycling
these networks would remain a problem. chain of its reactions, and another physical, due
It would seem fundamental that a living organ- to the frontiers produced by the membrane. In the
ism of any kind can be differentiated from cellular automata model of Varela (Varela
another. We know that the reproduction of a et al. 1974; McMullin and Varela 1994) illus-
second organism from a first is a central mecha- trated in Fig. 4, there are three types of particles
nism of life and can only operate if the “clone”
elaborates something to spatially distinguish
itself from its “original.” The best way of suc-
cessfully completing this individualization and to
be able to distinguish between these networks is
to revert to a spatial divide, which can only be
produced by some form of container capable of
circumscribing these networks in a given space.
Biochemists are well acquainted with an ideal
type of molecule, the raw material for these mem-
branes in the form of lipid/amphiphilic molecules
or fatty acids, the two extremities of which
behave in an antagonistic fashion – the first
hydrophilic, attracted to water, and the second
hydrophobic, repulsed by it. Quite naturally,
these molecules tend to assemble in a double
layer (placing the two opposing extremities oppo-
site to each other), formed by the molecules lin-
ing up and finally adopting the form of a sphere to
Artificial Life, Fig. 4 Simulation by means of cellular
protect the hydrophobic extremities from water.
automata of the autopoietic model originally proposed by
Like soap bubbles, these lipid spheres are semi- Varela. The minimal cell can easily be seen, together with
permeable and imprison the many chemical the catalysts and the substrates that it encapsulates
168 Artificial Life
capable of moving around a two-dimensional The whole, interactive “metabolism and mem-
surface: “substrates,” “catalysts,” and “links.” brane” prefigures a minimal elementary ▶ cell,
The working and updating rules of these cellular which already seems capable both of maintaining
automata go as follows: itself and detaching itself from its environment
and from cells similar to it. It is at this stage on the
• If two substrates are near a catalyst, they dis- way to establishing a better and more exact char-
appear to create one single link where one of acterization of life that the definition given by
the two was located. Luigi Luisi (2002) takes on its full meaning
• If two links are near each other, they link up (restating the idea of autopoiesis in more biolog-
and attach themselves to each other. Once ical terms). “Life is a system which can be
attached, these links become immobile. self-maintaining by using external energy and
• Each link is only allowed to attach itself to two nutritional sources to the production of its inter-
other links at the most. This allows the links to nal constituents. This system is spatially
form chains and to be able to make up a closed circumscribed by a semipermeable membrane of
membrane. its composition.” In the footsteps of Varela, con-
• The substrates can diffuse through the sidering life impossible without a way for
links and their attachments, while the individualization and compartmentalization, the
catalysts and the other links cannot. We can constitution of the membrane by simple self-
therefore understand how the process of the organization or self-assembly processes of bipo-
cogeneration comes about. The membranes lar molecules (hydrophilic and hydrophobic) has
shut in the catalysts and the links, become a very popular field of artificial life. It is
which in turn support the membrane by indeed rather simple to reproduce this phenome-
being essential to its formation and non in software (as illustrated in Fig. 5). You
regeneration. need water molecules that just randomly move,
• The reactions creating the links are reversible, in blue in the figure. You need two kinds of
as the links can recreate the two original sub- submolecules (call then A and B), which when
strates (and thus cause the membrane to dete- they meet – through the only authorized additive
riorate), but at a lower speed. When this
happens, the attachment between the links
also disappears.
Continuous updating and execution of these

rules produces minimal versions of reactive sys-
tems, physically closed and confined by means of
a membrane, which is itself produced by the
reactive system. For Varela and the others fol-
lowing him, this turns out to be an essential stage
in the road to life. Running the software, many
difficulties are encountered such as the simple
attainment of a closed cell on account of the
many more possibilities for the membranes to
unfold in a straight way. Only software simula- Artificial Life, Fig. 5 Simulation of a minimal cell based
tions can interconnect the physical compartment on A-B (A is hydrophobic and B hydrophilic) and water
played by the membrane with the generating molecules. All molecules move in reaction to repulsive
forces of different intensities and thermal agitations. The
metabolism and further show how far from obvi-
pink dots are the A; the gray dots are the B that do connect
ous it is for these two systems to mutually sustain to give A-B (represented in red and black) by a simple
each other. chemical reaction. The blue dots are the water molecules
Artificial Life 169
chemical reaction – form an A-B molecule (A is a cell to a computer and a genome to a code, we
hydrophobic and B hydrophilic) whose two poles need to explain how the computer itself was able
are connected by a small string. You need also to to be created out of this code. Let us follow the A
adjust the degree of repulsion between A and reasoning of this genius step by step, as it is the
water and between B and water, the strength of perfect illustration of an “artificial life” type of
the string of the A-B molecule, and the random approach: no material realization but just pure
component (akin to the thermal noise) to add on functions or rules. Through a sequence of purely
each of the intermolecular forces. Nevertheless, functional questions and showing an almost com-
the final outcome turns out to be rather robust. plete ignorance of actual biology, his reasoning
The bilayer of B-A/A-B molecules will very nat- led to a logical solution, the content of which
urally and spontaneously form just as for real retraces astonishingly closely those lessons we
cells. have since learned about the way biology func-
Again, as for Varela’s minimal cell, the clo- tions. Von Neumann begins from the principle
sure turns out to be quite delicate to obtain. One that a universal constructor C must exist, which,
very simple way to obtain it is to locate the source based on the plan of some kind of machine PM
of A submolecules (the pink dots in the figure) in (P the plan, M the machine), must be capable of
a singular point, so that the closed membrane will constructing the machine MP. This idea may be
simply surround that source, the circular shape simply translated by C(PM) = MP. The question
being the local minimal of the mechanical energy of self-replication that is then raised is, “Is this
connecting all A-B together. Like in Varela’s universal constructor capable of constructing
model, and somewhat paradoxically, the source itself?” In order to do so, it must, following the
needs to be circumscribed by the membrane for example of other construction products, have a
that same membrane to close on itself. However, plan of what it wants to construct; in this specific
in contrast with this autopoietic model, once in case, it is the constructor’s plan PC. The problem
place, the membrane cannot deteriorate, and thus, is then expressed as follows: can C(PC) give C
no further internal chemistry is required to endog- (PC) in order for there to be a perfect replication
enously produce what would be needed to fix of the original? Von Neumann therefore realized
it. Ultimately, this membrane should exhibit that the question at issue is that of the fate of the
some selective channeling in and channeling out construction plan, because if the constructor con-
(akin, for some authors (Luisi 2002), to a very structs itself, it has to add the plan itself to the
primitive form of cognition) providing its internal product of the construction. Von Neumann pro-
metabolism with the right nutrients and the right posed then allotting two tasks to the universal
evacuating way out so as to facilitate the cell’s constructor: constructing the machine according
self-maintenance. These two software models to the given plan and thus adding the original plan
raise interesting questions for the biologists like: to this construction. The constructor’s new for-
how are the molecular parts of the membrane mula then becomes C(PM) = MP(PM). If the con-
generated (endogenously or exogenously) and is structor applies itself to its own plan, this time the
this cogeneration of the membrane and the inter- replication will be perfect: CP(PC) = CP(PC).
nal metabolism the signature of minimal life? The fascinating aspect of Von Neumann’s
solution is that it anticipated the two essential
Self-Replication of This Elementary Cell functions that, as we have since discovered, are
Self-replication, or the ability of a system to the main attributions of the protein tools consti-
produce a copy of itself on its own, is one of the tuting the cell: constructing and maintaining this
essential characteristics that has most intrigued cell and also duplicating the code in order for this
and impassioned disciples of artificial life, begin- construction to be able to prolong itself for fur-
ning with John Von Neumann. Biology, and in ther generations. Starting with the ▶ DNA, the
particular this faculty of self-replication, fasci- whole ▶ protein machinery first of all builds the
nated Von Neumann. For if we want to compare cell and then, by an additional procedure,
170 Artificial Life
duplicates this same DNA. Von Neumann did not including those involved in the formation of
stop at duplication, because, at the same time, he membranes. This would also be the case provided
imagined how this same machinery could evolve all the reactions of the metabolism turn out to be
and become gradually more complex as a result autocatalytic, entailing the exponential growth in
of random mutations taking place while the plan the concentration of all its molecular elements
recopies itself. Von Neumann gave also a cellular (including again the membrane constituents).
automata solution of the problem in which each The membrane and the elements that it captures
cell of the automata possessed 5 neighbors and begin to grow (as illustrated in Fig. 7) until they
29 states, and around 200,000 cells were neces- reach the fatal point where the balance is upset.
sary for the phenomenon of self-replication to This is followed by the production of a new cell
take place. Many years later, Chris Langton produced by and from the old one. When the new
(Langton 1984, 1989), the organizer of the first one comes, it quickly grows fast enough to catch
conference on artificial life in 1989, proposed an up with the “generator” and “nursing” cell, as a
extremely simplified version of this (8 states, but chemical network is capable of some degree of
219 rules remain necessary), although it still fol- self-regeneration due to its intrinsic stability;
lows the pattern mapped out by Von Neumann. each molecule looks around for another that it
This automaton, shown in Fig. 6, incessantly can couple up to. This reconstitutes the natural
reproduces a little motif shaped as a loop. chain reaction of the whole. The new membrane
For many biologists, as opposed to Varela, and the new chemical network reconstitute on
Luisi, Ganti, and Maynard-Smith, life is not sim- their own by helping each other. Again, obtaining
ply indissociable from but also essentially reduc- such duplication is far from obvious since, any
ible to this capacity for self-replication. cell being intrinsically stable, only a thermal but
Nevertheless, they still need to explain how life quite unnatural agitation would do the job.
can actually reproduce without an entire preexis- Rather than this elementary form of chemical
tent metabolic chemical machinery. Departing self-replication coupled to the physical self-
from the elementary cell introduced in the pre- replication induced by the growth and division
ceding section and in the interest of an unbroken of the membrane, life has opted for a more
narrative, let us imagine a simpler scenario lead- sophisticated physicochemical version of it,
ing to self-replication. The closed circuit of more promising for the evolution to come: self-
chemical reactions could be destabilized by replication by the interposing of an “information
some kind of disturbance, causing a growth in template.” Each element of the template can only
concentration of some of its constituents, couple itself with one complementary element.
The new elements will as a whole naturally
reconstitute the template they were attracted to,
causing then the replication of the entire tem-
plate. In biology, it is the extraordinarily
emblematic double helix of DNA that acts as a
template, shouldering the major role in the his-
tory of life – that of the first known replicator. Our
elementary cell must now be internally equipped
with this information template. Since the 1950s,
Timor Ganti (2003) proposed the first minimum
mathematical system, named “chemoton,”
represented in the Fig. 8. This is the first abstract
computational protocell that we know,
constructed by Ganti as the original ancestor of
Artificial Life, Fig. 6 Langton’s self-replicating cellular living organisms. It possesses three autocatalytic
automata chemically linked subsystems: a metabolic
Artificial Life 171
Artificial Life, Fig. 7 The elementary minimal cell of network together with the membrane enclosing it. A lot
Fig. 5 in a process of self-replication induced by the of random thermal noise is here indispensable to destabi-
growing and the division of the chemical metabolic lize the initial cell
network, a membrane, and an information tem- Genetic Coding and Evolution by Mutation,
plate responsible for scheduling and regulating Recombination, and Selection
self-replication. All three grow exponentially In the information template introduced in the last
until they are able to reproduce, and they depend section, each letter constituting it contributes to
on each other for their existence and stability. The the code of a functional component essential to
metabolism feeds the membrane and the tem- the cell and designed on the basis of that code – a
plate, the membrane concentrates the metabo- protein. As soon as he hears anyone talking about
lites, and the template mechanism dictates the code, the software specialist has, quite legiti-
reproduction of the whole. The triad ensemble is mately, to put his head in through the window,
indeed capable of a whole synchronous self- because it is to him and him alone that we in fact
replication and tries to computationally answer owe the metaphor of the genetic code. Since
questions about the three subsystems and their ▶ Darwin and thereafter throughout all evolu-
interdependency, such as “how does the self- tionary science, we have a good idea of what the
replication of the template automatically accom- last chapter of the history of life is. Doubtless
pany the self-replication of the whole.” This com- what has stimulated most developments in “arti-
plex software object, the “chemoton,” has also ficial life” (primarily from the point of view of
become the topic of many software developments engineering) is the fact that the genetic code can
and experimentations and is emblematic of arti- evolve through ▶ mutation and sexual crossing
ficial life at its best. between the old machines, evolving so as to
172 Artificial Life
produce new machines that are more and more to show how beneficial this idea is for the
efficient. Over the past 20 years, many of those research and the automated discovery of sophis-
developments into artificial life have been eager ticated solutions to complex problems. As illus-
trated in Fig. 9, this research can take place
through a succession of mutations and recombi-
y nations operating at the level of the code, with the
best solutions proposed being preserved in the
x
next generation in order to be used for a new
Tm+k
cycle of these same operations. The brute force
of the computer is used to its full effect.
X A2
y
These are the same genetic algorithms that
A1 A1 Dawkins used in his Darwinian crusade, when
1 A3
pVn
he developed his biomorphs. It should also be
A5 V′ stressed that another element in Dawkins’ pro-
A4 pVnVn
2 gram is that, when it is finally evaluated, the
T′ R phenotype is not directly obtained from the geno-
pVnV′
type, as would be the case for classical optimiza-
tion in a real or combinatory space. In his work,
T
the biomorphs are the product of a recursive
sophisticated program, which is carried out
Tm 3 Tm+1 starting from a given ▶ genotype to give a
▶ phenotype. A great “semantic distance” is
Tm maintained between these genotypes and pheno-
types, which reflect the long process of cell con-
Artificial Life, Fig. 8 The schematic representation of
struction from the genetic code and the need for a
Ganti’s chemoton. One can easily see the three autocata-
lytic subsystems: the metabolism, the membrane, and the sophisticated metabolism building the machine
information template, chemically coupled out of the code. In brief, this constant program,
Artificial Life, GA Flowchart

Fig. 9 Illustration of the
genetic algorithms: the
recurrent iterated sequence Create initial design population
of selection, mutation, and
recombination easily leads
to an interesting solution of λ = 9000
a complex optimization λ = 5000 Evaluate obj. function of designs
λ = 1000
problem
old
Select and Reproduce
(Create new designs)
Generation
new
Next
Replace designs of the old

population with new designs
Stop?
Artificial Life 173
able to interpret the evolving genotype, is much ambiguously interpreted but commonly accepted
more important for the complexity of the final facts, and, when detailed at most, be able to
outcome than the genotype itself. Similarly, a predict experimental measurements. A
very prized derivative of these algorithms is The second partner, the engineer, is vigor-
genetic programming (Koza 1992), where the ously encouraged to use the computer for what
individuals now to be optimized are software it is best at doing – this infinite possibility of trial
codes. and error. There is a perfect synergy, where both
participants complement each other ideally: the
Conclusions engineer must bow to the computer in terms of
Parallelism, functional emergence, and adapt- calculating power, but this is compensated for by
ability are the conditions necessary to allow his judgment. Genetic algorithms, ant colonies,
these new biologically inspired artifacts to neural networks, and reinforced learning have
emerge, to “face the world.” We are jumping enriched the engineer’s toolbox.
straight into the robotics branch of artificial life Finally, for the philosopher, for each attempt at
(Brooks 1991). The interfacing with the real a definition of life, artificial life makes a real
world required by these robots needs a parallel attempt to achieve a computerized version in con-
information reception mechanism, because the formity with this definition. For the skeptic,
environment subjects it to a constant bombard- unhappy with this computerized “lining,” the
ment of stimuli. They have to learn to organize question now becomes how to refine his definition,
and master this avalanche falling on their percep- to complete it, or to renounce the possibility that
tions. They have to learn to build their own con- there is no definition that cannot be computerized.
cepts, fed and stimulated by this environment, The other possibility, doubtlessly more logical but
which, in turn, allows them to master it. The more difficult for many philosophers to accept,
conceptual high-level cognitive processes are would be that life poses no problem for a computer
born out of motor-sensory interactions and serve snapshot since it is computational at its roots.
to support them. Cognitive systems extend at new
levels what the minimal cell in the primitive soup
does, with a flow of matter and energy crossing
See Also
straight through, maintaining itself by selectively
integrating this influx to form a closed reactor
▶ Autocatalysis
network and the membrane enclosing it.
▶ Bioinformatics
My conclusions are addressed to the three
▶ Biological Networks
partners: the biologist, the engineer, and the phi-
▶ Cellular Automata
losopher. To the first, the outcomes of artificial
▶ Code
life consist in bringing out what the computer and
▶ Complexity
biology share intimately: an elementary way of
▶ Emergence of Life
working at the ultimate lowest level, but, which
▶ Genetic Algorithms
by the brute force of parallelism and incessantly
▶ Life
repeated iterations, can make unknown and
▶ Membrane
sophisticated phenomena to emerge at higher
▶ Self-Replication
levels. The qualitative aspect of these simulations
can give them new roles in the vast scientific
register: use it for education, illustrate biological References and Further Reading
principles that are already understood, open up
possible experiences of thought, play and replay Barabasi L-A (2002) Linked. The new science of net-
works. Perseus, Cambridge
multiple biological scenarios very quickly, titil- Bersini H (2004) Whatever emerges should be intrinsi-
late the imagination by on-screen representa- cally useful. In: Proceedings of artificial life,
tions, call into question some of the vol 9. MIT Press, Cambridge, pp 226–231
174 Artificial Meteorite
Billoud B (2010) Origins of life: computing and simula- Shapiro R (2007) A simpler origin for life. Sci Am
tion approaches. In: Gargaud M, Lopez-Garcia P, 296:46–53
Martin H (eds) Origin and evolution of life: an astro- Turing AM (1952) The chemical basis of morphogenesis.
biology perspective. Cambridge University Press Philos Trans R Soc Lond B 237:37–72; Also in
(Chapter 5) Saunders PT (ed) (1992) The collected works of
Brooks R (1991) Elephants don’t play chess. In: Maes A. M. Turing: morphogenesis. North-Holland,
P (ed) Designing autonomous agents. MIT Press, Cam- Amsterdam
bridge, MA Varela FR, Maturana HR, Uribe R (1974) Autopoiesis: the
Dawkins R (1986) The blind watchmaker. WW Norton, organisation of living systems, its characterization and
New York. ISBN 0-393-31570-3 a model. Biosystems 5:187–196
De Duve C (2002) Life evolving: molecules, mind, and
meaning. Oxford University Press, Oxford
Dyson F (1999) Origins of life, 2nd edn. Cambridge Uni-
versity Press, Cambridge
Fontana W (1992) Algorithmic chemistry. In: Langton
CG, Farmer JD, Rasmussen S, Taylor C (eds) Artificial Artificial Meteorite
life II: a proceedings volume in the SFI studies in the
sciences of complexity, vol 10. Addison-Wesley,
Reading ▶ STONE
Ganti T (2003) The principles of life. Oxford University
Press, Oxford
Goldberg DE (1989) Genetic algorithms in search, opti-
mization, and machine learning, 1st edn. Addison-
Wesley Professional, Reading
Kauffman S (1993) The origins of order: self-organization ASA
and selection in evolution. Oxford University Press,
Oxford Michel Viso
Kauffman S (1995) At home in the universe. The search
for the laws of self-organisation and complexity. CNES/DSP/SME, Vétérinaire/DVM,
Oxford University Press, New York Astro/Exobiology, Paris Cedex 1, France
Koza J (1992) Genetic programming. MIT Press,
Cambridge
Langton CG (1984) Self-reproduction in cellular autom-
ata. Phys D 10:135–144 Synonyms
Langton CG (ed) (1989) Artificial life I. Addison-Wesley,
Reading Aeronautics and Space Agency of FFG; Agentur
Lenaerts T, Bersini H (2009) A synthon approach to arti- f€ur Luft- und Raumfahrt der FFG; Austrian Space
ficial chemistry. Artif Life 15(1):89–103
Lovelock J (2000) Gaia: a new look at life on earth. Agency, Austria
Oxford University Press, Oxford
Luisi PL (2002) Some open questions about the origin of
life. In: Fundamentals of life. Elsevier, Paris,
pp 287–301. ISBN 2-84299-303-9 Definition
Maynard Smith J, Szathmary E (1999) The origins of life:
from the birth of life to the origin of language. Oxford The FFG’s Aeronautics and Space Agency
University Press, Oxford
(ASA) is the gateway to the international aero-
McMullin B, Varela FR (1994) Rediscovering computa-
tional autopoiesis. In: Husband P, Harvey I (eds) Pro- space industry for Austria’s industry and science
ceedings of the fourth European conference on sectors and aims to strengthen their international
artificial life. MIT Press, Cambridge, p 38 standing in these key technologies. The agency
Meinhardt H (1998) The algorithmic beauty of sea shells,
supports the participation of Austrian researchers
2nd edn. Springer, Heidelberg
Nagel T (1974) What is it like to be a bat? Philos Rev in international and bilateral aerospace collabo-
83:435–450; Repr. Mortal questions. Cambridge Uni- rations and fosters the creation and development
versity Press, New York, pp 165–180 of international networks. It implements Austrian
Plasson R, Kondepudi DK, Bersini H, Commeyras A,
aeronautical space policy and represents
Asakura K (2007) Emergence of homochirality in
far-from-equilibrium systems: mechanisms and role Austria’s interests in international aeronautical
in prebiotic chemistry. Chirality 19:589–600 and space organizations.
ASI 175
The FFG’s main focus is on managing the

contributions of the Republic of Austria to the Aseptic Process
programs of the ▶ European Space Agency A
(ESA), and FFG is responsible for the Catharine A. Conley
management of the Austrian Space Applications NASA Headquarters, Washington, DC, USA
Programme (ASAP), a bottom-up program
targeted to space science, technology,
space technology transfer, direct applications Definition
of space technology, and international
cooperation. Any operation which is carried out under condi-
Bilateral cooperative projects were under- tions that minimize the potential for contamina-
taken in particular with the former Soviet tion by ▶ microorganisms.
Union, such as the development of Austrian
instruments for space probes and missions.
These projects include, for example, the two
Venus probes, Venera 13 and 14 (1981–1982), ASI
the Vega 1 and 2 (1984–1986) missions to
Halley’s Comet, and the PHOBOS Mars Michel Viso
probes (1988–1989). The highlight of bilateral CNES/DSP/SME, Vétérinaire/DVM,
cooperation with the former Soviet Union Astro/Exobiology, Paris Cedex 1, France
was the AUSTROMIR-91 mission – the flight
of the first Austrian cosmonaut, Franz Viehböck,
to the MIR space station. Other bilateral projects Synonyms
were, and still are, run in partnership with Nor-
way, Sweden, France, Switzerland, and Agenzia Spaziale Italiana; Italian Space Agency
Germany.
Among other activities, FFG is organizing
annually since 1975, the well-known Definition
Alpbach Summer School that has a long
tradition in providing in-depth teaching on The Italian Space Agency was established in
aspects of space science and space technology 1988 to coordinate all of Italy’s efforts and
with the aim of advancing the training and investments in the space sector that had begun
working experience of European graduates, post- in the 1960s. Today, ASI has a key role at the
graduate students, young scientists, and European level where Italy is the third contribut-
engineers. ing country to the ▶ European Space Agency.
Number of employees of the Aeronautics and Italy is directly involved in major European
Space Agency of FFG is ten in 2010. and international programs. It provides several
elements for the International Space Station like
the multipurpose logistic module (MPLM) used
History to transfer cargo with the US space shuttle, Nodes
2 and 3, and is participating within ESA in the
The Space Research Institute of the Austrian European Automated Transfer Vehicle activities.
Academy of Sciences was founded in 1970, and ASI selected five astronauts flying either through
the Austrian Space Agency in 1972 (which had bilateral cooperation or through ESA. Franco
been merged into FFG in 2004). Austria has been Malerba was the first Italian in space in 1992
participating in ESA programs since 1975 and onboard the STS 46 flight. Italy is involved also
became a full member in 1987. in many missions dedicated to planetology,
176 Asparagine
astronomy, and exploration. It is playing a major O

role in the Cassini-Huygens mission, the
▶ ExoMars ESA mission, and many others.
Italy is taking also a major share (65 %) in the O
medium launcher program (Vega rocket) of ESA. OH
Beyond the headquarters in Rome, ASI has
three bases and one center. “Luigi Broglio”
Space Centre of Malindi, Kenya, was used to NH2 NH2
launch US Scout rockets with Italian satellites
from oceanic platforms (the marine segment) up Asparagine, Fig. 1 Structural formula of asparagine
to 1988. Nowadays this base (the ground seg-
ment) is dedicated to receive data from satellites
group (-CONH2) in its side chain, and it is easily
and launchers.
hydrolyzed to give ▶ aspartic acid, another pro-
A stratospheric balloon launch base is located
tein amino acid (Asp), and ammonia. In the
on a former airport in Trapani (since 1975). This
hydrolysis of proteins, both aspartic acid and
base is, in particular, actively involved in trans-
asparagine are determined as aspartic acid
Mediterranean flights of research balloons.
(noted Asx as the origin cannot be determined
In Matera, in collaboration with several insti-
unambiguously). It is classified as a neutral
tutions, ASI opened in 1983 a Space Geodesy
amino acid with an isoelectric point (pI) of 5.41.
Center dedicated to this discipline. Now this
base is diverting its activities welcoming some
technical activities required for robotic
See Also
exploration.
Finally, an ASI Science Data Center (ASDC)
▶ Amino Acid
was established in September 2000 for the man-
▶ Aspartic Acid
agement and analysis of scientific data collected
▶ Protein
by scientific satellites. This center is located in
the ESA facility (European Space Research
Institute-ESRIN) in Frascati, which is dedicated
to the Earth observation.
Aspartic Acid
Kensei Kobayashi
Asparagine Yokohama National University, Tokiwadai,
Kensei Kobayashi
Hodogayaku, Yokohama, Japan Definition
Aspartic acid, shown in Fig. 1, is one of the

Definition 20 protein ▶ amino acids. Its three-letter symbol
is Asp and one letter symbol is D. It is a
Asparagine, shown in Fig. 1, is one of the monoaminodicarboxylic acid, and it is classified
20 ▶ protein ▶ amino acids. Its three-letter sym- as an acidic amino acid. Aspartic acid has the
bol is Asn and one-letter symbol is N. It has a lowest isoelectric point (pI) 2.77 of the protein
molecular weight of 132.12. It has an amide amino acids. It is among the five amino acids that
Association Constant 177
O
Assimilative Metabolism
O
OH
A
Juli Peretó
OH NH2 Evolutiva, Universitat de València, València,
Spain
Aspartic Acid, Fig. 1 Structural formula of aspartic acid
were detected in Miller’s electric discharge

experiment in 1953 and is found in extracts Definition
from carbonaceous chondrites. Since it has two
carboxyl groups (a- and b-carboxyl group), it can Assimilative ▶ metabolism is the process by
make both a- and b-peptide bonds with other which an inorganic compound (NO 2
3 , SO4 ,
amino acids. In biosynthesis of ▶ proteins, only CO2) is reduced for use as a cell nutrient source.
a-peptide bonds are formed. As the ▶ racemiza- Assimilative metabolism is conceptually differ-
tion of aspartic acid is relatively rapid, the D/L ent from the ▶ reduction reactions that take place
ratio of aspartic acid can be used for dating bio- when the same inorganic compounds are used as
logical materials such as bone. electron acceptors to obtain energy by ▶ anaero-
bic respiration (▶ dissimilative metabolism).
There are important differences between both
See Also types of ▶ metabolism. In the assimilative metab-
olism, only enough of the compound is reduced to
▶ Amino Acid satisfy the needs for cell growth, and the products
▶ Miller, Stanley are normally converted into cell material, while
▶ Protein in the dissimilative metabolism, a large amount
▶ Racemization of ▶ electron acceptor must be reduced to guar-
anty the generation of sufficient energy and the
product is excreted into the environment.
Assay
See Also
Catharine A. Conley
▶ Anaerobic Respiration
▶ Dissimilative Metabolism
▶ Electron Acceptor
Definition
▶ Metabolism, Prebiotic
▶ Reduction
In ▶ planetary protection, an assay is the suite of
▶ Sulfate Reducers
actions performed during the integration of a
spacecraft or an instrument to collect and mea-
sure the biological contamination using a speci-
fied procedure, in order to estimate the number or
types of ▶ microorganisms associated with an Association Constant
item of interest (exposed surface, material, envi-
ronment, etc.). ▶ Affinity Constant
178 Asteroid
category of dwarf planets. Since about 1850

Asteroid the discovery rate has increased dramatically,
especially in recent years, leading to the
Alan W. Harris current tally (June 2013) of over 360,000 num-
DLR, Institute of Planetary Research, Berlin, bered asteroids. An asteroid is assigned a perma-
Germany nent designation, i.e., a sequential number, once
its orbit has become accurately established
through a sufficient number of astrometric
Keywords observations.
Asteroids and comets are considered to be
Dynamical and physical properties; Taxonomy; remnant bodies from the epoch of planet forma-
Mineralogy; Impact hazard; Mitigation tion. Planet embryos, roughly lunar-sized bodies,
formed in the protoplanetary disk about 4.5 bil-
lion years ago via the accretion of dust grains and
Synonyms collisions with smaller bodies (kilometer-sized
planetesimals). A number of planet embryos
Minor planet; Planetoid; Small Solar System succeeded in developing into the planets we
body observe today; the growth of other planet
embryos and planetesimals was terminated by
catastrophic collisions or a lack of material in
Definition their orbital zones to accrete. Most asteroids are
thought to be the fragments of bodies that formed
An asteroid is an irregularly shaped rocky body in the inner Solar System and were subsequently
orbiting the Sun that does not qualify as a planet broken up in collisions. Comets and related icy
or a dwarf planet under the International Astro- bodies are thought to have accreted in the cold,
nomical Union’s (IAU) definitions of those terms outer regions of the protoplanetary disk where
introduced in 2006. In contrast to planets and volatile materials, such as water and carbon diox-
dwarf planets, asteroids do not have sufficient ide, were abundant as ices.
mass for their self-gravity to overcome rigid Most numbered asteroids are in the main aster-
body forces and assume a hydrostatic equilibrium oid belt between the orbits of Mars and Jupiter.
(nearly round) shape. In contrast to comets, aster- The existence of the main belt is thought to be due
oids are inert bodies that do not display a coma of to the collisional fragmentation of remnant planet
gas and dust (although a few objects originally embryos and planetesimals that were prevented
classed as asteroids have subsequently been from accreting into planets by the gravitational
found to display cometary activity). Very small perturbations of the nearby massive planet
objects with a size of less than about 10 m are Jupiter.
normally referred to as meteoroids. Main-belt asteroids consist largely of silicates
and metals and come in all shapes and sizes up to
about 1,000 km in diameter. Table 1 lists physical
Overview data for the first ten asteroids discovered. Large
asteroids with diameters of several hundred km
The first asteroid, 1 Ceres, was discovered in tend to be roughly ellipsoidal, but smaller objects
1801 by the Italian astronomer Giuseppe Piazzi, generally have very irregular shapes. Asteroid
quickly followed in succeeding years by the dis- surfaces appear to consist of loose dust mixed
covery of 2 Pallas, 3 Juno, and 4 Vesta. Ironi- with gravel and boulders (regolith), whereby
cally, Ceres is now classed as a dwarf planet there is evidence that the regolith of km-sized
under IAU Resolution B5 of 2006, and Pallas bodies is coarser and less dusty than that of
and Vesta are candidates for transfer to this new large main-belt asteroids.
Asteroid 179
Asteroid, Table 1 The first ten numbered asteroids

Number Discovery: year, Taxonomic class Bulk density Rotation
and name site, discoverer Diameter (km) (see below) [4] (g cm3) [1] period (h) [5]
A
1 Ceres 1801, Palermo, 949 11 [1] G, C 2.12 0.04 9.074
G. Piazzi
2 Pallas 1802, Bremen, 533 6 [1] B 2.71 0.11 7.813
H. W. Olbers
3 Juno 1804, Lilienthal, 234 11 [2] S – 7.210
K. Harding
4 Vesta 1807, Bremen, 529 10 [1] V 3.44 0.12 5.342
H. W. Olbers
5 Astraea 1845, Driesen, 119 7 [2] S – 16.80
K. L. Hencke
6 Hebe 1847, Driesen, 185 3 [2] S – 7.274
K. L. Hencke
7 Iris 1847, London, 200 10 [2] S – 7.139
J. R. Hind
8 Flora 1847, London, 136 3 [2] S – 12.80
J. R. Hind
9 Metis 1848, Markree, 172 13[3] S – 5.079
A. Graham
10 Hygiea 1849, Naples, A. de 407 7 [2] C 2.76 1.2 27.62
Gasparis
References: [1] Britt et al. (2002); [2] Tedesco et al. (2002); [3] M€
uller and Barnes (2007); [4] Bus and Binzel (2002);
[5] Harris et al. (2008)
Asteroid Dynamical Groupings cloud of dust associated with the ecliptic plane
Asteroids are classified dynamically according to and give rise to the zodiacal light). In most cases
their orbital elements (semimajor axis, period, the family members have very similar
inclination, eccentricity, etc.). The most signifi- compositions.
cant grouping of asteroids is the main belt, Other major dynamical groups are described
between about 2.0 and 3.5 astronomical units below and listed in Table 2.
(AU, the mean Sun-Earth distance) from the Jupiter Trojans are asteroids that are trapped in
Sun. The main belt is populated by millions of dynamically stable zones 60 ahead of and
asteroids, some 360,000 of which have been behind Jupiter in its orbit. The stable zones are
assigned sequential numbers to date. associated with the L4 and L5 Lagrangian points
A number of asteroid families exist in the main of Jupiter’s orbit. There are some 5,900 known
belt. Family members have very similar dynam- Jupiter Trojans orbiting between about 5.0 and
ical characteristics and may be fragments from 5.4 AU from the Sun.
relatively recent (relative to the history of the Trans-Neptunian Objects (TNOs) are very dis-
Solar System) collisions. For example, there are tant, presumably icy, bodies with semimajor axes
large families associated with the main-belt aster- larger than 30 AU. Most TNOs are classed as
oids 4 Vesta, 8 Flora, and 10 Hygiea (Table 1). In asteroids according to the formal definition of
these cases the families presumably arose as the an asteroid given above, but in terms of their
result of cratering events that gave rise to ejecta physical characteristics, they may have more in
from the surfaces of the large asteroids. In other common with comets. The first TNO was discov-
cases precursor asteroids were apparently ered in August 1992; some 1,500 have been dis-
completely broken up. In both scenarios the result covered since.
is a family of fragments with similar orbits (and The Centaurs have orbits between that of Jupi-
dust particles that probably contribute to the ter and the TNOs. Centaurs are possibly objects
180 Asteroid
Asteroid, Table 2 Selected asteroid dynamical groups

Semimajor axis Approx.
Dynamical category [AU] no. knowna Notes
Trans-Neptunian objects >30 1,500
(TNOs)b
Centaursb 5.2–30 130 Possibly TNOs whose orbits have been
perturbed by Neptune
Jupiter Trojans 5.05–5.4 5,900 Associated with the Lagrangian points of
Jupiter’s orbit
Main belt 2.0–3.5 6.1 105 3.6 105 numbered asteroids
Amorsc >1 3,741 1.017 < perihelion 1.3 AU
Apollosc
1 5,373 Perihelion 1.017 AU
Atensc <1 774 Aphelion
0.983 AU
Inner-Earth objectsc <1 12 Aphelion < 0.983 AU
a
The listed approximate numbers of known objects are valid as of June 2013; these numbers increase rapidly with time as
new objects are discovered. For more details and updates, see the Minor Planet Center web site http://www.cfa.harvard.
edu/iau/mpc.html
b
TNOs and Centaurs are distant icy bodies that may have much in common with comet nuclei
c
The orbital characteristics of the different classes of NEAs are defined with respect to the Earth’s perihelion (0.983 AU)
and aphelion (1.017 AU) distances
that originated as TNOs but due to perturbations (Fujiwara et al. 2006), have provided a wealth of
by Neptune are now in orbits that bring them data on two very different NEAs in recent years
closer to the Sun. The number of known Centaurs (Figs. 1 and 2).
is currently around 130. A few Centaurs have
been observed to display comae and are classed The Taxonomic Classification and Mineralogy
as both asteroids and comets. TNOs and Centaurs of Asteroids
are of particular scientific interest because they Sunlight incident on the surface of an asteroid
have been subject to less thermal alteration and suffers absorption in particular wavelength
processing than main-belt asteroids and may con- bands depending on the minerals present;
tain well-preserved primordial material from the reflected light therefore carries a spectral signa-
epoch of formation of the Solar System. ture of the mineralogical composition of the
Near-Earth asteroids (NEAs) are asteroids that asteroid’s surface. Attempts have been made to
are thought to originate in the main belt but which classify asteroids according to details of the
now have highly evolved orbits with perihelion absorption features in their optical reflection
distances of less than 1.3 AU. NEAs are further spectra observed with astronomical telescopes.
categorized dynamically as Amors, Apollos, A number of classification schemes have been
Atens, or Inner-Earth objects (IEOs) according devised based on letters of the alphabet (e.g.,
to the semimajor axes, aphelion and perihelion Bus and Binzel 2002; Tholen and Barucci
distances of their orbits (Table 2). The orbits of 1989). For example, a very common spectral or
NEAs may evolve to intersect that of the Earth. taxonomic type is S, originally intended to sig-
Interest in the population of NEAs is focused nify a stony or “silicaceous” object. Another
mainly on the associated impact hazard (see common type is C, originally signifying carbon
below), but close approaches of NEAs to the rich or “carbonaceous.” Other letters, which are
Earth facilitate detailed telescope observations, in common use, are M for metallic and V for
including radar investigations, which provide Vesta-like. As the inventory of asteroid spectral
insight into the characteristics of asteroids in data grew, and more distinct spectral types were
general. Furthermore, two rendezvous missions, discovered, the taxonomic alphabet had to
NEAR-Shoemaker (Cheng 2002) and Hayabusa expand to incorporate more letters. Thanks to
Asteroid 181
the improved sensitivity of modern astronomical

instrumentation, it has become possible to iden-
tify subtle differences in spectral features within A
the classical taxonomic types, leading in some
cases to the addition of small letters after the
class letter to signify subclasses, e.g., Cb, Sq, etc.
Water and Organic Material in Asteroids

The mineralogical associations listed in Table 3
include hydrated (water-bearing) silicates and
organics. There is considerable evidence that
hydrated minerals are also present on a number
of M-type asteroids (Rivkin et al. 2000). Further-
more, water ice and organic material have been
detected on the surface of the main-belt C-type
asteroid 24 Themis (Rivkin and Emery 2010;
Campins et al. 2010). Some asteroids evidently
carry significant amounts of water and organic
Asteroid, Fig. 1 Near-Earth asteroid 433 Eros was the materials, a fact that may be relevant to questions
target of the NASA NEAR-Shoemaker spacecraft in
concerning the origin of water and life on Earth.
2000–2001. Eros is 34 km in length and of taxonomic
type S. The large number of impact craters indicates an It is widely believed that bodies similar to aster-
age of 1–2 Gyr (Credit: JHUAPL, NASA) oids and comets contributed to the early Earth’s
inventory of water and organics, but there is
currently no consensus as to when or how.
A point of current debate is whether Earth’s
water was provided primarily by asteroidal mate-
rial from the outer main belt at the time of for-
mation of the Earth or whether comets and
asteroids contributed a “veneer” of water-bearing
material at a later stage after the Earth had formed
and cooled. Measurements indicate that the
hydrogen isotopic ratio D/H in the few comets
for which such measurements have been made is
an order of magnitude higher than that of
protosolar material, but this significantly exceeds
the factor of 6 enrichment of the Earth’s oceans
(Lellouch et al. 2001; Drake and Righter 2002).
On the other hand, the D/H ratio of carbonaceous
chondritic asteroidal material is compatible with
that of the Earth’s oceans. Therefore, at face
Asteroid, Fig. 2 Near-Earth asteroid 25143 Itokawa was value, the D/H evidence appears to argue against
the target of the Japanese Hayabusa spacecraft in a dominant post-formation water contribution
2005–2006. Itokawa is 535 m in length and of type
from comets.
S. Itokawa, which lacks craters, is much younger than
Eros and may be an aggregate of components weakly Investigations of the composition of meteor-
bound by gravity or “rubble pile” (Courtesy of JAXA) ites, which are thought to originate in asteroids,
182 Asteroid
Asteroid, Table 3 Important asteroid taxonomic (spectral) classes

Associated geometric Approx. % of all
Class Probable mineralogy albedoa (pv) range classified asteroidsb
D, P Carbon, organic-rich silicates 0.03–0.06 2
C, B Carbon, organics, hydrated silicates 0.03–0.1 30
M Fe, Ni, enstatite 0.1–0.2 2
S Olivine, pyroxene, metals 0.1–0.3 40
Q Olivine, pyroxene, metals 0.2–0.5 1
V Pyroxene, feldspar 0.2–0.5 3
E Enstatite, other Fe-poor silicates 0.3–0.6 1
X Unknown (signifies otherwise unclassifiable 0.03–0.6 15
featureless spectrum)
a
The geometric albedo pv is the ratio of a body’s (V-band) brightness at zero-phase angle to the brightness of a perfectly
diffusing (Lambertian) disk with the same apparent size as the body
b
Estimated from the data of Bus and Binzel (2002) and Tholen and Barucci (1989). Note that discovery and observation
bias against objects with dark surfaces implies that the distribution of taxonomic classes in the known population may
not be representative of the entire asteroid population
indicate that carbonaceous chondritic asteroidal methods of determining asteroids masses (see
material contains up to 10 % by mass of water Britt et al. 2002 for a review), which all require
(Drake and Righter 2002; Morbidelli et al. 2000). measurement of the asteroid’s gravity field by
Therefore asteroidal material, in the form of means of, for example, a spacecraft, observations
planet embryos from the outer main belt incorpo- of the perturbations of the orbits of other asteroids
rated into the forming Earth (Morbidelli or Mars (applicable to large asteroids only), or
et al. 2000), may have contributed much of the observations of a satellite or companion asteroid
Earth’s water. On the basis of current knowledge, by means of precision optical or radar observa-
e.g., comparisons of isotope ratios and noble gas tions. Asteroid sizes can be determined from, for
ratios, it appears unlikely that either comets or example, spacecraft, thermal-infrared measure-
asteroids bombarding the Earth after the forma- ments (see Harris and Lagerros 2002 for a review
tion phase could have contributed significant and the section on asteroid physical properties
water (Drake and Righter 2002). below), radar observations (see Ostro
In the case of organic material, a post- et al. 2002, also Fig. 3), polarimetry,
formation source is required because it seems (e.g. Delbo’ et al. 2007), and occultation obser-
significant quantities of organics could not have vations (e.g. Dunham et al. 1990).
survived on the Earth until formation was com- Density estimates for just a few hundred aster-
plete and the Earth had cooled. The most plausi- oids are available to date (Carry 2012). An impor-
ble source of organic material is therefore the flux tant finding is that asteroid bulk densities tend to
of asteroids and comets that deposited material be significantly lower in general than expected
on the Earth after the formation phase. Some from measurements of meteorites and terrestrial
100 amino acids have been detected in meteorites analogues of asteroid material (2–8 g cm3). In
and it is widely believed that many comets and fact the bulk densities of some asteroids appear to
asteroids carry amino acids and other organic be similar to that of water (1 g cm3), implying
molecules. that these bodies must be highly porous. Such
results have increased speculation that some
The Densities and Structure of Asteroids asteroids, e.g., 25143 Itokawa, may be aggregates
Reliable estimates of density are difficult to of components of various sizes weakly bound by
obtain, since techniques for obtaining accurate gravity, or “rubble piles.” An asteroid that has
masses and sizes of asteroids are complex and been shattered by collisions with other objects
subject to large uncertainties. There are various may survive under the collective weak
Asteroid 183
Asteroid, Fig. 3 Radar images of binary near-Earth relative sizes or shapes of the components. The right-hand
asteroid 1999 KW4 (left-hand row of three frames) frame shows a computer model of the system in which the
received at the Goldstone 70-m antenna in May 2001. two components are seen to scale. The diameters of the
Each image has a resolution of 19 m/pixel and is a time primary and secondary bodies are 1.5 km and 0.5 km,
exposure spanning several hours, showing the motion of respectively (Images courtesy of L. A. M. Benner,
the secondary relative to the primary. Due to the complex- NASA/JPL)
ities of radar imaging, these three images do not show true
gravitational attraction of the resulting fragments given the potentially high relative velocities the
as a cohesionless, consolidated rubble pile; this effects in the case of a comet impact could be
idea is supported by the images of Itokawa much more devastating).
returned by the Hayabusa spacecraft (Fig. 2), The phenomenon of collisions in the history of
revealing a highly irregular object apparently our Solar System is very fundamental, having
consisting of separate component blocks of vari- played the major role in forming the planets we
ous shapes and sizes. observe today. Asteroids and comets may have
The rubble-pile idea leads to a natural expla- contributed to the delivery of water and organic
nation for the existence of the many binary aster- materials to the early Earth necessary for the
oid systems or asteroids with natural satellites development of life, but later impacts probably
(moons) that have been discovered (see Fig. 3 played a role in mass extinctions and they cur-
for an example). The spin rates of asteroids can rently pose a small but significant threat to the
be modified by dynamical phenomena, such as future of our civilization. Collisions of objects
the close approach to a planet, or the reflection with the Earth have taken place frequently over
and/or absorption and thermal reradiation of sun- geological history, and it is an irrefutable scien-
light. If a loosely bound rubble pile is spun up tific fact that major collisions with the Earth will
sufficiently, it can shed mass that may form a continue to occur at irregular, unpredictable
companion body or moon gravitationally bound intervals in the future.
to it. Recent computer simulations have demon- Collisions of asteroids and comets with the
strated the credibility of binary asteroid produc- Earth can have dramatic effects on the biosphere.
tion via the spin-up and rotational breakup of a A well-known example is the so-called
rubble pile (Walsh et al. 2008). Cretaceous–Tertiary (K-T) event 65 million
years ago, which is thought to have been caused
Near-Earth Asteroids and the Impact Hazard by the impact of an object with a diameter of
As a result of subtle thermal effects and the very 10–15 km, bringing about the extinction of not
strong gravitational field of Jupiter, small main- only the dinosaurs, but also more than 70 % of all
belt asteroids can drift into certain orbital zones species living at the time. The idea that the K-T
from which they may be ejected under the influ- mass extinctions observed in the paleontological
ence of Jupiter into the inner Solar System. As a record were caused by the impact of a near-Earth
result there exists a population of near-Earth asteroid was proposed by Alvarez et al. (1980),
asteroids, with orbits that can cross that of the and given a great deal of credibility by the dis-
Earth. Comets can also collide with the Earth but covery of a 65 million year old circular impact
the risk of a comet impact is thought to be much structure nearly 200 km in diameter centered near
lower than that of an NEA impact (although the town of Chicxulub on the Yucatan Peninsula,
184 Asteroid
Mexico (Pope et al. 1991). The impactor must The estimated impact frequency of NEAs on
have had a diameter of at least 10 km and may the Earth depends on size. The impact frequency
have hit the Earth with a velocity of some increases with decreasing size due to the size
15–25 km s1. The results of such an event distribution of the asteroid population: there are
would have included the deposition of billions many more small objects than large ones. The
of tons of dust and aerosols into the stratosphere impact risk to the Earth from NEAs based on a
that would have spread around the world, dim- recent assessment (Harris 2009) is summarized in
ming sunlight and cooling the surface for many Table 4.
months. The ensuing climatic effects could have
created a very stressful environment for life on Asteroid Physical Properties Relevant to the
the Earth’s surface for many years. Impact Hazard
While past impacts have probably altered the Accurate assessment of the impact hazard
evolutionary course of life on Earth, and paved depends on knowledge of the size distribution
the way for the dominance of mankind, we would and orbits of the NEA population. Which physi-
now rather not remain at the mercy of this natural cal parameters are most relevant for mitigation
process. Can we protect our civilization from the considerations? Preventing a collision with a
next major impact? Various initiatives are being NEA on course for the Earth would require either
taken by space agencies, including ESA and total destruction of the object, to the extent that
NASA, and research groups around the world to the resulting debris poses no hazard to the Earth
identify potential future impactors, investigate or, perhaps more realistically, deflecting it
their physical characteristics, and develop strate- slightly from its catastrophic course. In either
gies to mitigate against impacts. case accurate knowledge of the object’s mass
A number of observatories are operated spe- would be of prime importance. In order to
cifically to discover near-Earth objects and estab- mount an effective mission to destroy the object,
lish their orbits. The main currently active knowledge of its density, internal structure, and
asteroid search programs, such as the Catalina strength would also be required. Deflection of the
Sky Surveys, Lincoln Near-Earth Asteroid object from its course would require the applica-
Research (LINEAR), and Spacewatch (see tion of an impulse or continuous or periodic
Yeomans and Baalke 2009), are funded primarily thrust, the magnitude and positioning of which
by NASA and the US Air Force. Many observers, may depend on the mass and its distribution
including amateur astronomers, around the world throughout the (irregularly shaped) body, the sur-
contribute to the tracking of asteroids and submit face characteristics, and the spin vector,
their astrometric data to the Minor Planet Center depending on the strategy deployed. Mitigation
in Cambridge, Massachusetts, which acts as a planning takes on a higher level of complexity if
clearing house for asteroid position data and the Earth-threatening object is a rubble pile or
maintains databases of orbits. binary system.
As of June 2013 the number of known near- Since the observed brightness of an asteroid is
Earth objects is approaching 10,000, of which proportional to its surface albedo and cross-
some 860 have diameters of 1 km or more; the sectional area, a very rough size estimate can be
total number of the latter, including as yet obtained from the observed brightness and
undiscovered objects, is thought to be about knowledge of the asteroid’s orbit. However, the
1,000 (see below). A special term, “potentially albedo can have any value between 3 % and 60 %
hazardous asteroids” (PHAs), is reserved for (Table 3), so the assumption of a typical albedo
those with diameters above 120 m and orbits of, say, 15 %, can lead to errors in diameter and
that bring them within 0.05 AU of the Earth’s mass of a factor of 2 and 8, respectively. While
orbit. PHAs are large enough to survive passage current estimates of the overall impact hazard are
through the Earth’s atmosphere and cause exten- necessarily based on the assumption of a typical
sive damage on impact. or mean albedo for NEAs, more accurate
Asteroid 185
Asteroid, Table 4 Estimated frequency and effects of asteroid impacts on the Earth
Crater
Impactor size (m) Mean impact Energy released diameter Possible effects/
larger than interval (year) (megatons TNT) (km) comparable event
A
30 300 2 – Fireball, shock wave,
minor damage
50 2,000 10 1 Tunguska-type explosion
or small crater
100 10,000 80 2 Largest H-bomb detonation
200 40,000 600 4 Destruction on national
scale
500 200,000 10,000 10 Destruction on continental
scale
1,000 600,000 80,000 20 Many millions dead, global
effects
5,000 20 million 10 million 100 Billions dead, global
climate change
10,000 100 million 80 million 200 Extinction of human
civilization
The energy release estimates assume a density of 3,500 kg m3 (stony body) and an impact velocity of 20 km s1. The
given impact intervals are statistical: for example, the probability of a 100-m or larger object impacting in the next
100 years is 1 %, which is the same as the probability of such an object not impacting in 46,000 years
methods of size determination are clearly neces- nearly 12 months and observed a total of at least
sary for accurate hazard estimation and mitiga- 584 NEAs, of which more than 130 were new
tion purposes. discoveries.
Telescope observations in the thermal infrared An important result from thermal-infrared
combined with knowledge of an asteroid’s opti- observations is the apparent size dependence of
cal brightness offer a means of obtaining more the albedos of some types of NEA in the
accurate information on size and albedo (Harris 0.1–10 km size range, such that the mean albedo
and Lagerros 2002). Darker, low albedo, aster- appears to increase with decreasing size (Delbo’
oids are less reflective in the visible spectral et al. 2003; Harris 2006). As explained above, the
region and absorb more solar radiation; they are mean albedo of NEAs is important for current
therefore warmer and brighter in the thermal- estimates of their size distribution. If the mean
infrared spectral range. The opposite is true for albedo is higher, the diameter derived from a
high albedo asteroids. The two different physical particular observed brightness is smaller and the
relationships governing visible and thermal- overall impact hazard from NEAs is reduced
infrared brightness enable simultaneous solutions (Stuart and Binzel 2004). The impactor sizes in
for size and albedo to be obtained. While the Table 4 are based on a mean geometric albedo of
number of asteroids observed in the thermal 14 %, slightly larger than the value of 11 %
infrared is still only a very minor fraction of the adopted in earlier studies, which leads to some-
total known, this technique has provided the vast what longer mean impact intervals for each size
majority of size and albedo determinations to category.
date. For example, surveys of the sizes and albe-
dos of hundreds of NEAs have been carried out
by the NASA WISE (Wide-field Infrared Survey Future Directions
Explorer) and Spitzer infrared space telescopes.
WISE was launched to Earth orbit in December The rate of discovery of NEAs has increased
2009 carrying a 40-cm-diameter telescope and dramatically in recent years and is now seriously
infrared detectors. WISE surveyed the sky for outstripping the rate at which the population can
186 Asteroid
be physically characterized. The NEA population

is still largely unexplored. Telescope observa-
tions will continue to provide valuable informa-
tion about the NEA population as a whole, but
rendezvous missions are vital for probing the
detailed characteristics of individual objects. To
date only two NEAs have been visited by space-
craft (Figs. 1 and 2), but many more rendezvous
missions to NEAs of different taxonomic types
will be necessary before we can start to under-
stand the diverse compositions, structure, and
origins of NEAs and their relation to meteorites.
Mitigation of Hazardous Asteroids

At present there is no general agreement on the
most effective strategy to adopt in the case of a
predicted impact. In the case of an object with a
diameter below 100 m, the best course of action
may be to simply evacuate the region around the
predicted impact point (see Table 4), assuming
there would be sufficient advance warning (only a Asteroid, Fig. 4 Artist’s impression of a kinetic-
small fraction of the asteroids in this size cate- impactor spacecraft deployed to modify the orbit of a
near-Earth asteroid. The second spacecraft at the bottom
gory have been discovered to date). For objects of the picture is orbiting the asteroid or “hovering” near it
larger than 100 m, a number of mitigation strate- to observe the effects of the impact and monitor the altered
gies may be considered, depending on trajectory of the asteroid. The scenario depicted derives
circumstances. from the Don Quijote mission, studies of which were
commissioned by ESA in 2006. The European-led
The NASA report to Congress (2007) on the NEOShield project is currently carrying out detailed
surveying and deflection of near-Earth objects investigations of this and other promising deflection tech-
concluded that nuclear devices offer the most niques (Credit: ESA-AOES Medialab)
effective means of applying a deflecting force to
an asteroid. While they may offer the only feasi- produced in the impact (Fig. 4), predictions of
ble solution in desperate circumstances, e.g., in which would require some prior knowledge of the
the case of very little advanced warning, it is asteroid’s physical properties. To facilitate effec-
widely felt that the obvious political problems tive mission planning, the kinetic-impactor
associated with launching nuclear devices and approach would require an initial reconnaissance
testing them in space seriously compromise the mission to gather relevant physical data.
practicability of this technique. The NASA report Alternative approaches include the “gravity
concluded that the most effective nonnuclear tractor” and “space tug.” The gravity tractor
option is the kinetic impactor, which involves relies on the force of gravity between the target
applying an impulsive force to the asteroid by asteroid and a spacecraft hovering under power in
means of a large mass in the form of a spacecraft close proximity to gradually modify the aster-
accurately guided to the target at a high relative oid’s orbit. A significant advantage of the gravity
velocity. This technique was the subject of stud- tractor is that no contact with the target is
ies commissioned by ESA in 2006 of a dual- required. The principle of the space tug is similar,
spacecraft mission called Don Quijote (ESA but in this case, the spacecraft is physically
NEO Space mission Preparation 2006). The attached to the asteroid’s surface and prior
change of momentum of the target asteroid knowledge of surface characteristics would be
depends on the porosity and the amount of ejecta necessary for effective mission planning.
Asteroid 187
Excess emission from HD69830

0.20 Crystalline
Crystalline
olivine (forsterite) A
0.15 olivine Crystalline
olivine
Flux density (Jy)
Amorphous Crystalline
olivine Crystalline olivine
proxene Crystalline
olivine
0.05
0.00
−0.05
10 15 20 25 30 35
Wavelength (μm)
Scaled emission from Hale-Bopp

0.20 Crystalline
olivine (forsterite)
Crystalline
Flux density (Jy)
Amorphous olivine Crystalline

olivine Crystalline
0.10 olivine olivine
Crystalline Crystalline
proxene olivine
0.05
0.00
−0.05
10 15 20 25 30 35
Wavelength (μm)
Asteroid, Fig. 5 Comparison of features in the spectra of Hale Bopp, detailed analysis of the spectral signature of
the excess thermal emission from the star HD 69830 and the dust around HD69830 indicates that it probably orig-
the comet Hale Bopp. The similarity of the spectra inates from the breakup of asteroids (Lisse et al. 2007).
strongly suggests that the dust in the debris disk around The figure is taken from Beichman et al. (2005) and is
HD 69830 has a very similar composition to dust in the reproduced by permission of the AAS
Solar System. Despite the similarities to the spectrum of
The gravity tractor and space tug are “slow-push” and the USA, was established in 2012 to investi-
approaches that require sustained, reliable pro- gate deflection techniques in detail. The project,
pulsion and sophisticated autonomous control called NEOShield (www.neoshield.net), aims to
systems to achieve the required amount of deflec- provide solutions to scientific and technical
tion, but may be promising techniques in cases in issues that will enable the feasibility of promising
which there is sufficient advance warning (e.g., mitigation options to be demonstrated in the
decades), the target is relatively small, and/or a future via test missions. Research into the
very slight, precise deflection is required to pre- mitigation-relevant physical properties of
vent an impact on the Earth. NEOs, including laboratory experiments and
A unique research program funded by the associated modeling, is aimed at supporting tech-
European Commission and involving 13 partner nological development work leading to the
organizations in 6 countries, including Russia detailed design of demonstration missions.
188 Asteroid
Asteroids in Extrasolar Planetary Systems ▶ Chondrite

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Stuart JS, Binzel RP (2004) Bias-corrected population, ▶ planetesimals and planetary embryos
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190 Asteroseismology
cause them to be ground down into an ever-

increasing number of smaller bodies. Asthenosphere
Daniele L. Pinti
See Also Geodynamics, Université du Québec à Montréal,
▶ Asteroid
▶ Jupiter
▶ Mars
Definition
▶ Planet
▶ Planetesimals
The asthenosphere is the viscous mechanically
▶ Solar System, Inner
weak-ductile region of the upper ▶ mantle of
the Earth. It lies below the ▶ lithosphere, at
depths between 100 and 200 km, and it is
involved in the plate tectonic movements and
Asteroseismology isostatic adjustments. The heat is transmitted by
convection and the thickness and viscosity of
Patrick Eggenberger this layer controls the stress applied at the base
Geneva Observatory, University of Geneva, of the lithosphere. This, in turn, influences
Geneva, Switzerland the global tectonic style of the Earth and of
any other terrestrial planet. The viscosity and
the convective regime of the asthenosphere
Synonyms has also a control on the degassing history of a
terrestrial planet and the evolution of its
Stellar seismology atmosphere.
Definition
See Also
Asteroseismology is the study of stellar oscilla-
▶ Lithosphere, Planetary
tions aiming at determining the internal structure
▶ Magma
and global properties of stars. The oscillation
▶ Mantle
patterns observed on the stellar surface result
▶ Plate Tectonics
from waves that propagate into the interior of
the star. The observation of these oscillations,
done either by monitoring the variation of the
star’s brightness or of its radial velocity, offers
therefore a unique means to probe the otherwise Astrobiology
unobservable internal layers of stars and to help
us in our understanding of the complex physical William M. Irvine
processes at work in stellar interiors. University of Massachusetts, Amherst, MA, USA
See Also Synonyms
▶ Stellar Pulsation Bioastronomy; Exobiology

Astrometric Orbit 191
Definition Moreover, many questions relevant to astrobi-

ology have in fact been asked by humanity
This encyclopedia defines astrobiology as the throughout recorded history and certainly before A
study of the origin, evolution, and distribution that. In consequence, the Encyclopedia includes a
of life in the universe (see Preface). Such section on History of Science, which includes
studies, or some part of them, have also been theories on the origin and evolution of life and
referred to as exobiology or bioastronomy on the possible multiplicity of habitable worlds
(the term employed by the International from classical Greek and Roman times, from the
Astronomical Union), and at the present time, Islamic world, from ancient Asia, from
these terms can be considered synonymous. pre-Columbian America, and from the modern
Although it has been stated that astrobiology world.
is a science without a subject (since life Lastly, what is the future of life, at least on
outside the Earth remains undiscovered), in Earth? Entries concerning ▶ artificial life
fact there is a vast and growing volume of are included which address some of these
research in fields included in the present questions.
encyclopedia.
See Also
Overview
▶ Enceladus
Thus, although the physical and chemical pro-
▶ Europa
cesses that led to the origin of life on Earth are
▶ Evolution, Biological
still far from understood, they clearly involve,
▶ Mars
inter alia, the nucleosynthesis of chemical ele-
▶ Planetary Theories and Cosmology, Islamic
ments including carbon that are essential for life;
Theories
the formation and evolution of the Sun and
▶ planets, including the delivery of water and
▶ Solar System, Inner
organic material to the ▶ Earth; prebiotic chemi-
▶ Solar System, Outer
cal processes; and the nature and evolution of the
▶ Titan
early Earth’s atmosphere, crust, and interior. Fur-
▶ Water in the Solar System
thermore, the study of the evolution of life and
▶ Water in the Universe
ultimately the development of ▶ intelligence on
our planet requires research, e.g., in biochemistry,
biophysics, microbiology (including life in envi-
ronments that seem extreme by human standards),
and ▶ genetics. Astrometric Orbit
There is also the question of whether life exists
elsewhere in the solar system or at other locations David W. Latham1 and Nader Haghighipour2
1
in the universe. These issues are addressed in the Harvard-Smithsonian Center for Astrophysics,
Encyclopedia of Astrobiology through Cambridge, MA, USA
2
entries characterizing various bodies in the solar Institute for Astronomy, University of Hawaii-
system (particularly the existence of water, the Manoa, Honolulu, Hawaii, HI, USA
universal solvent of terrestrial life), current knowl-
edge concerning planets around other stars
(extrasolar or exoplanets), how life might be iden- Definition
tified beyond the Earth (biosignatures or ▶ bio-
markers), and space missions to investigate these When two bodies are in ▶ Keplerian orbits
topics. around a common center of mass, the apparent
192 Astrometric Planets
motion projected onto the plane of the sky defines

an astrometric orbit. In some cases, such as the Astrometric Planets
famous example of Sirius A and its white dwarf
companion Sirius B, both objects can be spatially G. Fritz Benedict1 and Nader Haghighipour2
1
resolved, and their absolute positions and McDonald Observatory, The University of
motions can be determined. This allows the Texas, Austin, TX, USA
2
ratio of the two masses to be determined, which Institute for Astronomy, University of Hawaii-
can be combined with the total mass, if the dis- Manoa, Honolulu, Hawaii, HI, USA
tance is known, to derive individual masses.
More often, only a single photocenter can be
resolved, and only motions relative to nearby Keywords
stars can be determined. In this case an orbital
solution requires additional information about the Astrometry; Coplanar orbits; Orbital inclination;
relative brightness of the components. In the case Perturbation; Planetary mass; HST FGS
of a planetary companion, the contribution of the
planet to the total light can almost always be
neglected, and the mass of the planet can be Definition
estimated from the observed wobble of the parent
star if the mass of the star and its distance can be Astrometry is a discipline of astronomy that is
estimated. In common with a spectroscopic orbit concerned with the measurements of stellar posi-
determined from radial velocities along the line tions. As applied to exoplanets, a series of astro-
of sight, an astrometric orbit also yields the metric measurements over time is analyzed to
▶ period, ▶ eccentricity, and time of obtain an estimate of the size of the reflex motion
▶ periastron passage. Astrometry has the of the host star caused by the gravitational pull of
advantage over radial velocity of measuring its unseen planet. The planet and its host star orbit
motions in two dimensions on the plane of the a common center of mass. Given an estimate of
sky instead of just one motion along the line of the mass of the host star, normally from the mass-
sight, and this allows the orbital inclination to be luminosity relation, one can determine the mass
determined. Thus, astrometric orbits can elimi- of the planet. Astrometry also provides the orbital
nate the ambiguity of the unknown orbital incli- inclination of exoplanets. The knowledge of the
nation and can yield actual masses under certain orbital inclination of a planet will enable us to
circumstances, for example, when applied to determine the true value of its mass (see “▶ Incli-
minimum masses determined by spectroscopic nation (Astronomy)”) and can also be used to
orbits. constrain models of planet formation.
History
See Also
The history of astrometric planets is littered with
▶ Astrometric Planets past failures. A notable example was the reported
▶ Eccentricity discovery of a planetary system associated with
▶ Exoplanets, Discovery Barnard’s Star by van de Kamp (1969) which was
▶ Inclination (Astronomy) subsequently disproven through reanalysis of the
▶ Keplerian Orbits data by Ianna (1995) and re-observation of the
▶ Periastron same system by Benedict et al. (1999). Astrome-
▶ Period try has yet to discover an independently verified
Astrometric Planets 193
exoplanet; although as a technique to further axis = 0.8 mas for the outermost companion,
characterize the companions to stars discovered HD 38529 c, motivated us to obtain astrometric
by precision Doppler measurements (radial observations with mas precision using HST FGS, A
velocities), it has enjoyed some recent successes. to determine the orbital inclination and thus the
Future successes (both discovery and characteri- true mass of this planet. These astrometric data
zation) are expected from the astrometric satellite span 3.25 years of the complete 5.85-year period.
Gaia.
Basic Methodology
Overview
Our approach depends on the discovery and
We describe some of the astrometric planet orbital characterization results from radial veloc-
results obtained using one of the Fine Guidance ity studies. Specifically, the period, eccentricity,
Sensors (FGS) aboard the Hubble Space Tele- time of periastron passage, and position angle of
scope (HST). The techniques employed are illus- periastron (P, e, T, o) of the perturbing body are
trative of both past astrometric approaches and initially known. As can be seen from the
future astrometric investigations. Exoplanetary Table below, most of our target systems have
astrometry relates the time-varying position of a periods far in excess of 2 years. The data from
target to positions of stars defining a reference the astrometry and radial velocity are combined
frame. To obtain the orbital elements of the per- through the constraint (Pourbaix and Jorissen
turbation due to the companion, one must first 2000), a sin i/pabs = P K(1 e2)1/2/
characterize motions extrinsic to the system: par- (2p 4.7405), where quantities derived only
allax and proper motion (defined below). Anyone from astrometry (parallax, pabs; apparent
attempting to measure a perturbation will find semimajor axis of the elliptical perturbation, a;
(to either their delight or horror) that the Earth and inclination, i) appear on the left, and quanti-
orbits the ▶ barycenter of the Solar System. The ties derivable from both the period P and eccen-
corresponding apparent motion of the host star tricity e, or from radial velocity only (the RV
evidences itself as an ellipse, which for nearby amplitude of the primary K), appear on the
stars can exceed 100 milli-arcsec (mas) in size. right. In most cases such as the one here, given
Added to that parallactic motion is the apparent the fractional orbital coverage of the HD 38529 c
motion of the host star across the line of sight due perturbation afforded by the astrometry, all the
to the transverse component of the star’s space quantities on the right-hand side are dominated
velocity relative to our Sun. This proper motion by the radial velocities (Table 1).
often exceeds 50 mas/year. We use the determi- Before analysis, all astrometric measurements
nation of the mass of the planetary companion from the FGS are corrected for optical field angle
HD 38529 c (Benedict et al. 2010) as an example distortion. Like the part of an iceberg below the
of the methodology. The star HD 38529 (= HIP water, this correction, while typically “out of
27253 = HR 1988 = PLX 1320) hosts two sight” (McArthur et al. 2003), is an essential
known companions b and c, discovered by high- part of our process. Actual FGS distortions with
precision ▶ radial velocity (RV) monitoring (see amplitudes in excess of 1 arcsec are reduced to
Wright et al. 2009). Previously published mini- 1 mas or less over the FGS field of regard.
mum masses of HD 38529b and c were 0.85 and We are able to relate the position of the bright
13.1 Jupiter masses, respectively. The latter lies host star to the far fainter reference frame stars
just above the official definition of the transition through the use of a neutral density filter. The
between planets and brown dwarfs. A predicted astrometric reference frame for HD 38529 con-
minimum perturbation of the semimajor sists of four stars. Any prior knowledge
194 Astrometric Planets
Astrometric Planets, Table 1 HST FGS astrometric results. Fe/H measures the abundance of iron relative to hydrogen
on a logarithmic scale normalized to the Sun. Sp.T. is the spectral type of the host star
Companion M*(MO) [Fe/H] Sp.T. d(pc) ecc M(MJup) a (mas) inc( ) P(d)
GJ 876 b 0.32 0.122 M4 V 4.7 0.1 1.9 0.5 0.25 84 6 61
55 Cnc d 1.21 +0.32 G8 V 12.5 0.33 4.9 1.1 1.9 53 7 4,517
e Eri b 0.83 0.03 K2 V 3.2 0.7 1.6 0.2 1.9 30 4 2,502
HD 33636 B 1.02 0.13 G0 V 28.1 0.48 142 11 14.2 4.0 0.1 2,117
HD 136118 b 1.24 0.01 F9 V 52.3 0.35 42 15 1.5 163.1 3 1,191
HD 38529 c 1.48 +0.27 G4 IV 40.0 0.36 17.6 1.4 1.1 48.3 3.7 2,136
u And c 1.31 +0.15 F8 V 13.5 0.25 14 4 0.62 81 240.94
u And d 1.31 +0.15 F8 V 13.5 0.32 10 2 1.4 24 1 1281.51
HD 128311 C 0.84 0.02 K0 V 16.5 0.16 3.8 0.7 0.46 56 14 921.5d
concerning these four stars eventually enters our unsuccessful, but did yield a planetary mass for
modeling as observations with errors, which component c (McArthur et al. 2014).
helps improve the values of the parallax and
proper motion for the prime target, HD 38529.
Distances to the reference stars are estimated Applications
from (BVIJHK) photometry and stellar classifi-
cation spectroscopy. Of particular value are inde- These techniques are presently being applied to
pendently measured proper motions from the determine masses for 14 Her b, HD 47536 b, and
UCAC3 catalog (Zacharias et al. 2010). All HD 168443 c. There now exist sufficient FGS
these periodic and nonperiodic motions must be astrometric measurements to attempt to deter-
removed as accurately and precisely as possible mine orbital inclinations for b and c, HD
to obtain the orbital inclination, i, and perturba- 202206 b and c, m Ara b and c, and g Cep A and
tion size, a, caused by HD 38529 c. b. The latter two host stars may be too bright for
Gaia. If the degree of orbital coplanarity can be
established for those systems, astronomers will
Key Research Findings have valuable data to probe the architectural
uniqueness (or not) of our own Solar System,
Results from HST astrometry are presented in the complementing analogous results from the recent
Table above. HD 38529 c is clearly more massive Kepler transit discoveries and the study of
than 13 Jupiter masses, the threshold above ▶ radial velocity planets.
which deuterium fuses to helium. This threshold
has been adopted by the International Astronom-
ical Union as the official transition between Future Directions
planets and brown dwarfs.
Other results: McArthur et al. (2010) deter- Major progress will result from two improve-
mined the inclinations of components c and d in ments to past practice as exemplified by HST
the u Andromeda system, indicating FGS astrometry. First, future missions will
non-coplanarity. Whether this architecture is a employ global (as opposed to local) reference
result of the migration of component b (with frames, removing the correction to absolute par-
Msini = 0.69 Mjup) now in a P = 4.6-day allax required when using a local reference
orbit, or the past passage of the M star compan- frame. Second, the astrometric precision will
ion, u Andromeda B, is as yet unknown (Barnes improve by two orders of magnitude, from milli-
et al. 2011). A recent attempt to determine the second of arc to micro-arcseconds (mas). Future
degree of coplanarity for HD 128311 was success is expected from Gaia, an astrometric
Astrometry 195
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▶ European Space Agency (ESA). The expected Whitmore B (eds) The 2002 HST calibration work-
shop. STScI, Baltimore, p 373
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should permit the determination of thousands of the υ andromedae system with data from the hubble
exoplanet masses and the degree of orbital copla- space telescope and hobby-eberly telescope.
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orbiting a nearby Sunlike star at a star-planet hipparcos transit data and intermediate astrometric
data of spectroscopic binaries. I. Ba, CH and Tc-poor
separation allowing surface liquid water, remains S stars. A&AS 145:161
a difficult goal, one requiring 1 mas per observa- Standish EM Jr (1990) The observational basis for JPL’s
tion precision. DE 200, the planetary ephemerides of the astronomical
almanac. Astron Astrophys 233:252
van de Kamp P (1969) Alternate dynamical analysis of
Barnard’s star. Astron J 74:757
Wright JT et al (2009) Ten new and updated multiplanet
See Also systems and a survey of exoplanetary systems.
Astrophys J 693:1084
Zacharias N et al (2010) The third US naval observatory
▶ Aeronautics and Space Agency of FFG CCD astrograph catalog (UCAC3). Astron J 139:2184
▶ Barycenter
▶ Doppler Shift
▶ European Space Agency
▶ Exoplanets, Discovery
▶ Gaia Mission Astrometry
▶ Radial-Velocity Planets
▶ Spectral Type François Mignard
CNRS, Observatoire de la Côte d’Azur,
University of Nice Sophia-Antipolis, Nice,
France
Barnes R et al (2011) Origin and dynamics of the mutually
inclined orbits of υ andromedae c and d. Astrophys
J 726:71 Keywords
Benedict GF et al (1999) Interferometric astrometry of
proxima centauri and barnard’s star using HUBBLE
SPACE TELESCOPE fine guidance sensor 3: detec- Coordinate systems; Fundamental astronomy;
tion limits for substellar companions. Astron Gaia; Hipparcos; Reference frame
J 118:1086
Benedict GF et al (2010) The mass of HD 38529c from
hubble space telescope astrometry and high-precision
radial velocities. Astron J 139:1844 Synonyms
Casertano S et al (2008) Double-blind test program for
astrometric planet detection with Gaia. Astron Positional astronomy
Astrophys 482:699
Heintz WD (1978) Double stars. Reidel, Dordrecht
Ianna PA (1995) Barnard’s star: twenty years of McCor-
mick observations. Astrophys Space Sci 223:161 Definition
Lindegren L et al (2008) The Gaia mission: science, orga-
nization and present status. In: Jin WJ, Platais I, Astrometry is that part of astronomy dealing with
Perryman MAC (eds) A giant step: from milli- to
micro-arcsecond astrometry. Proceedings of the IAU the determination of the position, distance, and
Symposim 248. Cambridge University Press, Cam- motion of celestial bodies and by extension their
bridge, pp 217–223 size and shape. This is by far the oldest branch of
196 Astrometry
astronomy, and until the mid-1800s the word did system astrometry or to determine the relative
not even exist. It was only coined at that time to motion within a binary system. Its accuracy
make the distinction from the new field of can reach 1 mas (0.001 arc sec) in relative
astrophysics. positions.
• Wide-angle astrometry was the basis for the
construction of the fundamental catalogues
Overview and their extension to fainter magnitudes.
The classical instruments were the meridian
Taken as a broad subject, astrometry covers the circles and the astrolabes. Stars over the whole
definition and realization of the astronomical celestial sphere were tied together, thanks to
▶ coordinate systems and the construction and the rotation of the Earth or by overlapping
maintenance of positional star catalogues, but frames in photographic or ▶ CCD astrometry.
also the techniques for the computation of astro- The accuracy, including zonal errors, is of the
nomical events, like eclipses, passages of inner order of 20 mas for the bright stars (visual
planets across the Sun’s disk, orbits of ▶ binary ▶ magnitude V <8) and 50–70 mas at around
stars, stellar ▶ occultation by the Moon, or the V = 15.
minor planets. Classical astrometry is undergoing • Space astrometry is the only way to do abso-
a true revolution with the access to radio interfer- lute measurements over the whole sky with a
ometry and above all the possibility to carry out single instrument able to connect widely sep-
accurate measurements from space, thanks to the arated directions (>60 ) and perform repeated
Hipparcos mission (1989–1996) and in the near observations over several years. This method
future the Gaia mission. The main benefit of leads to an absolute catalogue of positions
doing astrometry in space is the measurement of and ▶ proper motions, independent of any
absolute ▶ parallaxes of stars, that is to say their preexisting reference frame. It is aligned to
distances, just from the geometric method with- the radio frame constructed on extragalactic
out any assumptions on the physics of the sources sources by using sources observed by
or on the absorption of the light during its journey both techniques. Accuracy was 1 mas with
to the telescope. ▶ Hipparcos (120,000 sources) and
Given the fundamental nature of its investiga- should be close to 0.025 mas at V = 15 with
tions, it is fair to state that astrometric data, lead- ▶ Gaia and several hundred million
ing to the realization of an accessible ▶ reference sources. Radio ▶ interferometry performs
frame or giving the distances and the masses of also absolute astrometry of quasars on the
the stars, provide the foundation of stellar physics whole celestial sphere. The associated
and of the cosmic distance ladder. Stellar dis- catalogue is currently the basis of the astro-
tances are the key to determining stellar luminos- nomical reference frame (ICRF: International
ities and then for calibrating further methods to Celestial Reference Frame) with an accuracy
determine distances up to the most remote of about 0.1 mas over a few hundreds of
galaxies. sources.
Based on the tools employed and on the objec-
tives, astrometry can be further divided up into
three broad categories: See Also
• Small-angle astrometry, measuring the ▶ Coordinate Systems

position or the motion of stars in a very small ▶ Gaia Mission
field (less than 1 ) with respect to local refer- ▶ Hipparcos
ences. This was typical of astronomical pho- ▶ Magnitude
tography until the advent of charged coupled ▶ Parallax
device (▶ CCDs) and is still used for solar ▶ Proper Motion
Asymmetric Reaction, Absolute 197
References and Further Reading known as asymmetric induction. One example of

absolute asymmetric synthesis is the introduction
Green RM (1985) Spherical astronomy. Cambridge Uni- of optical activity into helicenes (twisted
versity Press, Cambridge A
polyaromatic hydrocarbons with six rings) stud-
Kovalevsky J (2002) Modern astrometry, 2nd edn, Astron-
omy and astrophysics library. Springer, Berlin ied by Kagan et al. (1974) in 1971, in which the
Kovalevsky J, Seidelman PK (2004) Fundamentals of optical yield was lower than 1 %. Enantiomeric
astrometry. Cambridge University Press, Cambridge enrichment of racemic compound such as
Murray CA (1983) Vectorial astrometry. Adam Hilger,
amino acids by irradiation with circularly polar-
Bristol
Walter H, Sovers OJ (2000) Astrometry of fundamental ized light (CPL) is one of the well-known exam-
catalogues, Astronomy and astrophysics library. ples of asymmetric decomposition. The first
Springer, Berlin experimental evidence of CPL-induced asym-
metric decomposition was reported by Kuhn
and Knopf (1930).
Various light sources such as lasers and syn-
Astronomical Unit chrotron radiation are now available as powerful
tools to study asymmetric reactions. Several stud-
▶ AU ies have examined the role of asymmetric reac-
tions on the origin of ▶ homochirality in
biological molecules: Flores et al. (1977),
Asymmetric Reaction, Absolute Nishino et al. (2001), and Meierhenrich
et al. (2005).
Kazumichi Nakagawa Polarized radiation from pulsars and interstel-
Graduate School of Human Development and lar masers has been known for several decades.
Environment, Kobe University, Nada, Natural sources of circularly polarized light were
Kobe, Japan found in space and reported by Bailey
et al. (1998). Mie scattering and synchrotron
radiation are thought to be the natural sources of
Keywords circularly polarized light.
Other mechanisms of absolute asymmetric
Asymmetric decomposition; Asymmetric synthe- reactions have been discussed by Bonner
sis; Circularly polarized light; Origin of et al. (1974), Bonner (2000), and Goldanskii
homochirality; Induction of chirality and Kuzmin (1989).
Definition
See Also
An absolute asymmetric chemical reaction is one
▶ Chirality
induced by asymmetric physical conditions such as
▶ Homochirality
circularly polarized light (CPL) instead of by
▶ Polarized Electron
chemical reagents or catalysts. They can be divided
▶ Polarized Light and Homochirality
into two categories: one is asymmetric synthesis
and the other is asymmetric decomposition.
Overview Bailey J, Chrysostomou A, Hough JH, Gledhill TM,

MacCall A, Clark S, Menard F, Tamura M (1998)
Circular polarization in star-formation regions: impli-
Asymmetric synthesis is driven by the introduc- cations for biomolecular homochirality. Science
tion of chirality into an achiral system, a process 281:672–674
198 Asymptotic Giant Branch Star
Bonner WA (2000) Parity violation and the evolution of (fusion) of the hydrogen and He shells. In the
biomolecular homochirality. Chirality 12:114–126 He shell, neutrons are released through (a,n)
Bonner WA, Kavasmaneck PR, Martin FS (1974) Asym-
metric adsorption of alanine by quartz. Science reactions and are captured on preexisting nuclei
186:143–144 of the Fe peak to produce heavier elements
Flores JJ, Bonner WA, Massey GA (1977) Asymmetric (s-process). The thermal pulses occur in time-
photolysis of (RS)-leucine with circularly polarized scales of 104–105 years. They mix material from
ultraviolet light. J Am Chem Soc 99:3622–3624
Goldanskii VI, Kuzmin VV (1989) Spontaneous breaking the burning shells to the convective envelope (the
of mirror symmetry in nature and the origin of life. Sov 3d dredge up), and they also induce heavy mass
Phys Usp 32:1–29, Asp Fiz Nauk 157:3–50 loss from the star (more than 50 % of its mass),
Kagan HB, Balavoine G, Moradpour A (1974) Can circu- creating a ▶ planetary nebula.
larly polarized light be used to obtain chiral com-
pounds of high optical purity? J Mol Evol 4:41–48
Kuhn W, Knopf E (1930) Darstellung optisch aktiver
Stoffe mit Hilfe von Licht. Z Phys Chem Abteil See Also
B 7:292–310
Meierhenrich UJ, Nahon L, Alcaraz C, Bredehöft JH,
Hoffmann SV, Barbier B, Brack A (2005) Asymmetric ▶ Hertzsprung-Russell Diagram
vacuum UV photolysis of the amino acid leucine in the ▶ Planetary Nebula
solid state. Angew Chem Int Ed 44:5630–5634 ▶ S-process
Nishino H, Nakamura A, Inoue Y (2001) Synchronous
enantiomeric enrichment of both reactant and product ▶ Stellar Evolution
by absolute asymmetric synthesis using circularly
polarized light. Part 1. Theoretical and experimental
verification of the asymmetric photoisomerization of
methyl norbornadiene-2-carboxylate to methyl
quadricyclane-1-carboxylate. J Chem Soc Perkin Atacama Desert
Trans 2:1693–1700
Richard Léveillé
Natural Resource Sciences, McGill University,
Asymptotic Giant Branch Star
St. Anne de Bellevue, Québec, Canada
Nikos Prantzos
Definition
Synonyms The Atacama is a hyperarid desert located in

northern Chile. It is considered to be one of the
AGB driest places on Earth with average annual rain-
fall of less than 1 mm in some locations. It is a
useful analogue to the arid surface of Mars and
Definition for analog studies of hydrology, mineralogy, and
microbiology. Features that may be analogous to
An asymptotic giant branch (AGB) star is a low- Mars also include caves, salt deposits, water-
or intermediate-mass star (of mass M < 8 M) at derived mudflows, and gully-type deposits.
a late evolutionary phase in its life, during which Field tests of prototype instruments and rovers
it appears as a red giant in the ▶ Hertzsprung- have also been conducted there.
Russell diagram. After the star has exhausted the
supply of He for fusion in its core, it draws energy
from He fusion in a shell around the inert carbon- See Also
oxygen core, in the early AGB (E-AGB) phase.
Later, the star enters the thermally pulsing ▶ Desiccation
(TP-AGB) phase, with intermittent burning ▶ Extreme Environment
Atmosphere Escape 199
▶ Extremophiles Overview
▶ Mars Analogue Sites
▶ Terrestrial Analog The thermal escape velocity is obtained by equat- A
ing initial kinetic energy to the gravitational
potential energy:
pffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi pffiffiffiffiffiffiffi
Atacama Large Millimeter Array 1=2 mv2 ¼ GMP m=r, v¼ 2GMP =r ¼ 2gr
▶ ALMA where m is the mass of the object (molecule in the

present case), v its speed, r its initial distance
from the center of the planet, MP the mass of the
planet, G the universal gravitational constant, and
Atacama Large Millimeter/ g the acceleration of gravity at distance r from the
Submillimeter Array planet’s center. In order to allow a molecule to
escape, enough energy must be delivered to the
▶ ALMA molecule to accelerate it to the escape velocity.
Since the kinetic energy of a molecule with mass
m is ½ mv2, light molecules like H2 will move
faster and hence escape more easily than heavier
molecules like N2, given an equal delivery of
Atmosphere Escape energy. Dissociation of a molecule like CO2 or
H2 into lighter individual components also aids
Ray Pierrehumbert escape. Using the formula for escape velocity, we
Department of the Geophysical Sciences, can define the escape energy of a molecule with
University of Chicago, Chicago, IL, USA mass m as mgr. For escape of N2 from altitudes
not too far from the Earth’s surface, this energy is
2.9 1018 J; for H2 the escape energy is only
Definition 0.2 1018 J.
Atmospheric escape may be categorized as
Atmospheric escape refers to the loss of a planet’s follows:
atmosphere to space. Broadly speaking, escape
may be energy limited or diffusion limited. For • Thermal (Jeans) escape – molecular speeds
the energy-limited case, permanently removing a generally obey the Maxwell-Boltzmann distri-
molecule from a planet’s atmosphere can be com- bution. Fast-moving species in the high-
pared to sending a rocket from Earth to space: one energy tail of the distribution may attain
must impart enough velocity to the object, and in enough energy to escape the atmosphere. The
the right direction, to allow the object to over- general thermal energy of the atmospheric gas
come the potential energy at the bottom of the ultimately comes from, e.g., absorbed stellar
gravitational well and still have enough kinetic radiation or from heat leaking out of the inte-
energy left over to allow the object to continue rior of the planet.
moving away. The study of atmospheric escape • Nonthermal escape mechanisms. Here, a “col-
in this case thus amounts to the study of the lisional process” energizes gas species above
various ways in which the necessary energy can the escape barrier. For example, “ion
be imparted to molecules to reach the escape exchange” involves charged species, e.g.,
velocity, which is the minimum velocity an hydrogen cations (H+), which are accelerated
object needs in order to escape to infinity, pro- in the planet’s magnetic field to high energies.
vided no drag forces intervene (Pierrehumbert If the H+ acquires an electron from a neigh-
2011). boring neutral species, it is converted into a
200 Atmosphere, Model 1D
neutral, high-energy atom (H*) which can mean atmosphere depend only on the altitude
escape the atmosphere. Nonthermal escape above the planetary surface, so that the depen-
can also involve interaction of, e.g., atmo- dence is one-dimensional (a 1D atmosphere). The
spheric species with the stellar wind (e.g., vertical structure of the atmosphere is determined
“sputtering” refers to the collisional ejection by the pressure, temperature, density, and chem-
of a neutral species from the atmosphere by ical composition with height above the surface.
incoming high-energy species). “Impact ero- The average atmospheric profile of the Earth
sion” refers to atmospheric escape driven by reproduces the flux detected if the Earth were
high-energy impacting material. seen from far away as an exoplanet, thus provid-
ing a disk-integrated view of our planet. The
assumption of an average atmospheric profile is
See also
very useful to explore a variety of effects on
exoplanet atmospheres.
▶ Absorption Cross Section
▶ Scale Height
See Also
References and Further Reading ▶ Atmosphere, Structure

▶ Grey Gas Model
Pierrehumbert RT (2011) Principles of planetary climate. ▶ Non-Grey Gas Model: Real Gas Atmospheres
Cambridge University Press, Cambridge
▶ Scale Height

Atmosphere, Model 1D
Chamberlain J (1987) Theory of planetary atmospheres.
Lisa Kaltenegger Academic, New York
Cornell University, Ithaca, NY, USA
Synonyms Atmosphere, Organic Synthesis
Model atmospheres Henderson James (Jim) Cleaves II

Definition Japan
The simplest atmosphere is gravitationally bound USA
with no vertical motions, i.e., in hydrostatic equi- Blue Marble Space Institute of Science,
librium and spherically symmetric. For terrestrial Washington, DC, USA
planets, solar radiation and the radiative proper- Center for Chemical Evolution, Georgia Institute
ties of the atmospheric components provide a of Technology, Atlanta, GA, USA
first-order description of the vertical thermal
structure. An atmosphere controlled by stellar
light can hardly be spherically symmetric; never- Definition
theless, it is very useful to think of a mean plan-
etary atmosphere, i.e., averaged over, e.g., It is widely thought that the origin of life on a
latitude and day-night. The properties of such a planet may depend on the availability of
Atmosphere, Structure 201
abiotically synthesized organic compounds. Var- composed of hydrogen and helium are thought
ious potential sources for these exist, including to form as a result of gas accretion by a growing
extraterrestrial delivery and submarine hydro- planet embedded in a ▶ protoplanetary disk. For A
thermal synthesis. However, one of the first low-mass planets or even Moon- to Mars-sized
suggested means of producing these compounds planetary embryos, a relatively small amount of
was via synthesis in the gas phase in planetary gas may be gravitationally captured into a prim-
atmospheres, as shown by the results of the now itive atmosphere. The atmospheres of full-size
famous experiment published by Stanley Miller terrestrial planets are then made up of a combi-
in 1953. It has now been amply demonstrated that nation of atmospheres from their constituent
the action of various forms of energy, including planetary embryos, as well as volatiles that may
shock waves, electricity, and ionizing and UV be degassed from the planetary interior or deliv-
radiation acting on various simulated planetary ered to the planet via impacts of small bodies
atmospheres can result in the synthesis of a vari- during final stages of accretion (▶ late veneer).
ety of organic compounds. In general, more In that case a large range of atmospheric compo-
reduced gas mixture atmospheres appear to give sitions is possible, and the atmospheres can con-
more favorable yields of organic compounds; sist of water and other volatile materials.
however, even relatively oxidized gas mixtures
still produce organics to some extent.
See Also
See Also ▶ Earth’s Atmosphere, Origin and Evolution of

▶ Late Veneer
▶ Electric Discharge ▶ Protoplanetary Disk
▶ Miller, Stanley
▶ Proton Irradiation
▶ UV Radiation
Atmosphere, Structure
John Lee Grenfell

Atmosphere, Primitive Envelope German Aerospace Center (DLR), Berlin,
Germany
Sean N. Raymond
Laboratoire d’Astrophysique de Bordeaux,
CNRS, Universite de Bordeaux, France Definition
Atmospheric structure usually refers to changes

Synonyms in physical quantities such as temperature, den-
sity etc. in the vertical. Atmospheres can be clas-
Primary atmosphere; Primitive atmosphere sified into regions (troposphere, stratosphere,
mesosphere etc.) depending upon the variation
of temperature in the vertical. In lower regions
Definition where convection is important, the decrease in
pressure with height leads to expansion hence
An atmosphere (or an atmospheric envelope) typ- adiabatic cooling. Warm air lies below cool air
ically refers to the gas that is gravitationally and vertical mixing is strong – this is the “tropo-
bound to a planet or another planetary object. In sphere” region. At higher altitudes, temperature
relation to planet formation, atmospheres may increase with altitude due to absorption of
202 Atmosphere, Temperature Inversion
some of the incoming UV energy, e.g., on Earth absorption of sunlight by molecules or aerosols
via ozone. At pressures lower than about 0.1 bar that are chemically produced. There exists a
(on Earth, Jupiter, Uranus) radiative transfer highly nonlinear positive feedback between
starts to dominate over convection. Cool air lies chemistry, radiation, and dynamics, such that
below warm air and vertical mixing is weak – this absorbers will remain longer, hence lead to a
is the “stratosphere” region. At higher still alti- larger thermal inversion, resulting in the trapping
tudes, temperature may revert back to a decrease of even more absorbers (Pierrehumbert 2011).
with altitude e.g., on Earth above about 50 km
where ozone concentrations are small – this is the
mesosphere region. Above the mesosphere lies Overview
the thermosphere, where temperature starts to
increase again in the vertical e.g., on Earth due In the troposphere, convection dominates and the
to oxygen absorption of high energy radiation. temperature decreases according to the so-called
Finally, above the thermosphere lies the wet lapse rate, about 6.5 K/km for Earth (which
exosphere – the uppermost atmosphere region includes the latent heat released by the conden-
before space. The bottom of the exosphere, the sation of water). Mixing is generally rapid in a
exobase, occurs where the atmospheric scale troposphere, as the warm, lighter air at the bottom
height equals the molecular mean free path. The tends to rise and the colder air at the top tends to
top of the exosphere can be defined to occur sink down. The thermal inversion in the strato-
where the stellar pressure equals the planetary sphere on Earth is due to absorption of ultraviolet
gravitational pull on a hydrogen atom. radiation by Ozone (O3) but can be generated by
other molecules on different planets. Above
Earth’s stratosphere where O3 is no longer
strongly produced, the temperature decreases
Atmosphere, Temperature Inversion with height due to cooling in the infrared, mainly
by CO2. In the outermost atmosphere, Earth’s hot
Lisa Kaltenegger thermosphere is related to absorption of extreme
Cornell University, Ithaca, NY, USA ultraviolet radiation by O2, N2, and O.
In contrast to the Earth, Mars has a thin and
Venus a thick atmosphere, which is reflected in
Keywords their surface temperatures. The tropospheric tem-
peratures of Mars and Venus follow their respec-
Atmosphere structure; Biomarkers; Habitability; tive lapse rates (atmospheric thermal gradient).
Habitable zone; Temperature inversion There are no well defined stratospheric inver-
sions, except during dust storms on Mars. The
thermospheres of both planets are relatively cold
Definition compared to Earth’s, because the major atmo-
spheric gas (CO2) is a very effective radiator of
A temperature inversion refers to a reversal in the thermal energy. In contrast, the primarily apolar
normal decrease of atmospheric temperature with constituents of Earth’s atmosphere (N2, O2) have
altitude in the lower portion (▶ troposphere) of a virtually no bands in the infrared and are poor
planetary atmosphere (▶ atmosphere, Structure). radiators.
Temperature inversions inhibit vertical motion,
because the warmer, less dense air is at a higher
altitude and tends to remain there. The net result See Also
of a temperature inversion is a long time constant
for exchange between a stratosphere and a tropo- ▶ Adiabatic Processes
sphere. It exists due to the heating produced by ▶ Albedo
Atmospheric Habitability 203
▶ Apolar Molecule gravitational field and for a terrestrial planet

▶ Atmosphere, Model 1D extends from the surface up to the exosphere.
▶ Atmosphere, Structure This entry discusses the critical importance of A
▶ Clouds atmospheres for habitability.
▶ Greenhouse Effect
▶ Latent Heat
▶ Scale Height Overview
“Habitable” is derived from the Latin habitabilis

meaning “to inhabit.” “Habitable” denotes envi-
Pierrehumbert RT (2011) Principles of planetary climate. ronmental conditions which could support life
Cambridge University Press, Cambridge (Steele et al. 2005) but does not necessarily
imply that life is present. In simple terms one
can imagine a house which is habitable but may
be unlived in. In its simplest terms habitability is
Atmosphere-Ocean General classically defined as the requirement for liquid
Circulation Model water. However, “conditions which could sup-
port life” can be rather more many and varied.
▶ AOGCM Broadly speaking, one can define three central
criteria, namely, the access to (i) energy, (ii) a
suitable solvent (e.g., water), and (iii) molecular
complexity (e.g., via carbon chain molecules).
Atmospheric Dusts Other factors such as the availability of nutrients
and protected conditions (e.g., shelter from UV,
▶ Aerosols cosmic rays, climate excesses, strong impacts,
etc.) also play a role. The distribution of habit-
ability in the Universe depends, therefore, on how
well we understand the limits and occurrences of
Atmospheric Habitability these various criteria and on how sensitively life
responds to them. In this entry we will focus on
John Lee Grenfell the central role for habitability played by plane-
German Aerospace Center (DLR), Berlin, tary atmospheres in and beyond the Solar System.
Germany Atmospheres are essential for habitable con-
ditions for several reasons. First, liquid water
(needed for habitability) is thermodynamically
Keywords unstable at low pressures. Second, (in addition
to a magnetosphere) atmospheres protect the
Habitable; Habitable zone; Life; Atmospheric planetary surface, e.g., from harmful radiation
and cosmic rays. Third, atmospheres transport
heat from warm to cool regions which helps to
Definition maintain stable conditions favored by life.
Fourth, atmospheres may interact with biological
“Habitability” is classically defined as the (past and geochemical processes via feedback cycles
or present) existence of surface conditions which such as the “carbonate-silicate” cycle which sta-
support life. This in turn leads to the requirement bilizes the (Earth’s) climate. Fifth, (on Earth) the
for the presence of liquid water which is needed atmosphere supplies CO2 for autotrophic life and
by all known life. “Atmosphere” refers to the O2 needed for aerobic respiration and hence
envelope of gas trapped within a planet’s directly supports higher life on our planet.
204 Atmospheric Habitability
Finally, atmospheric spectral signatures on rocky inner HZ boundary. Applying an atmospheric

exoplanets may provide indications of surface column model, Kopparapu et al. (2013) suggest
habitability and even biosignature signals. that using updated water absorption coefficients
and applying the same approach as in Kasting et
al. (1993) lead to the inner HZ moving out to
Key Research Findings 0.99 AU.
Regarding the outer HZ, here, CO2 clouds
Huang (1960) investigated the key requirements could form in the atmosphere although their radi-
a star should meet to maintain life in its habitable ative properties are challenging to estimate since,
zone (HZ). Dole (1964) introduced the “complex e.g., their shapes and sizes are not well defined.
life habitable zone (HZ)”– the region where a The “maximum greenhouse effect” is defined to
planet could have surface temperatures between occur at the furthest distance of planet to star for
0 and 30 C (for >10 % of its surface with an which a planetary surface temperature of 273 K
oxygen (O2)-rich atmosphere and <1.5 Earth can be held for a planet having a cloud-free, CO2
gravity). Early climate modeling by Hart (1978) atmosphere.
suggested a narrow HZ from 0.95 to 1.01 astro-
nomical units (AU) for a solar-type star. Atmospheric Evolution
Kasting et al. (1988) subsequently suggested The solar neighborhood includes some rather
that including negative (i.e., opposing the origi- young stars (e.g., Woolley et al. 1970) which
nal change) feedback mechanisms such as the could have planets resembling the early Earth in
carbonate-silicate cycle could significantly age. The Earth’s atmosphere has changed signif-
expand the HZ. Kasting et al. (1993) calculated icantly over geological time. The above has moti-
the HZ boundaries for different main sequence vated studying the early Earth in an exoplanetary
stars. context. For example, Grenfell et al. (2011)
investigated different epochs of hypothetical
Modern Solar System Earth-like exoplanets. The Sun’s net luminosity
The (classical) HZ in the Solar System lies some- was (20–30 %) weaker on the early Earth
where between Venus and Mars. There are two (~2.5–4.0 Gyrs ago) compared with the modern
HZ inner boundaries which may be defined. Earth. Although observations from ancient soils
Firstly, the “water loss limit” (~0.95 AU, Kasting (paleosols) suggest the presence of liquid water
et al. 1993) occurs where a planet would lose its on early Earth, numerous climate model studies
mass of ocean within its current lifetime due to (see, e.g., Kasting et al. 1988) imply a completely
atmospheric escape. Secondly, the “runaway frozen planet – the so-called Snowball Earth. The
greenhouse limit” (~0.84 AU, Kasting contradiction between paleosol data and model
et al. 1993) occurs where the planetary surface predictions is named “the faint young Sun prob-
temperature exceeds the critical point of water. lem.” Several possible solutions have been put
Atmospheric clouds if present could strongly forward, e.g., supplying additional greenhouse
influence the HZ boundaries. Kitzmann gases such as methane (CH4) or nitrous oxide
et al. (2010) investigated the radiative effects of (N2O) (Grenfell et al. 2011) to the early Earth
clouds upon atmospheric temperature for Earth- atmosphere. Goldblatt et al. (2009) suggested a
like planets orbiting F, G, and K stars. Results stronger greenhouse effect on early Earth due to
suggest that cloud type and altitude could signif- enhanced atmospheric N2 (considering up to
icantly impact surface temperature. The HZ three times the modern value). Greenhouse
limits, as estimated from numerical model stud- warming on early Earth is also expected to be
ies, also depend on, e.g., planetary mass, density, sensitive to surface pressure (P0). Som
and atmospheric mass and composition. et al. (2012) suggested that early Earth featured
Regarding the inner HZ, Abe et al. (2011) var- (P0 < 2) bars, based on fossilized mud
ied ocean mass and calculated the effect on the droplet data.
Early Venus and Mars and Gaidos (2011) suggested that collision-
How early Venus’ atmosphere evolved, whether/ induced greenhouse warming of high pressure
for how long the surface was habitable, and how (~40 bar) H2 protoatmospheres (which may be A
it finally diverged into its modern extreme state relevant for super-Earths) could also lead to a
are critical questions to address. On Mars, significant increase of the outward HZ
observed surface flow features suggest that liquid boundary – maybe to as far as 10 AU for solar-
water likely existed early in its history. Analo- like stars.
gous to the early Earth, there are difficulties for
numerical studies to reproduce sufficiently warm Waterworlds and Desertworlds
surface temperatures. The study by von Paris “Waterworld” here refers to a terrestrial exoplanet
et al. (2013) suggested that including pressure with complete ocean coverage, i.e., without
broadening (which enhanced the atmospheric exposed surface continents. “Desertworld” here
greenhouse effect) of N2 can help address this refers to a terrestrial exoplanet whose surface
issue. How the Martian atmosphere developed lacks large-scale oceans and vegetation. On
from possible habitable conditions to its waterworlds, it is currently unclear whether stabi-
modern-day state is challenging for interior, lizing climate cycles, i.e., analogous to the
atmospheric, and impact models. Clearly, under- carbonate-silicate cycle on Earth, could operate.
standing the developments of Venus and Mars More work on, e.g., seafloor carbonization and its
will provide useful insights for understanding dependence on, e.g., pressure, pH etc. are required
the HZ boundaries around different stars. (see, e.g., Wordsworth and Pierrehumbert 2013).
Regarding desertworlds, additional work estimat-
Beyond the Solar System ing, e.g., the dependence of ocean mass (e.g.,
Recent works have expanded the parameter range including the lower limit) on, e.g., plate tectonics
with the aim of determining which are the key and habitability would be useful.
factors affecting the atmospheric habitability of
Earth-like planets beyond the Solar System. Fac- Earth-Like Planets Orbiting M Dwarf Stars
tors investigated include, e.g., climate feedbacks, These are favored targets for exoplanet search
interactive atmospheric climate chemistry (e.g., missions. Such objects are already starting to be
Segura et al. 2003; Grenfell et al. 2007), radiative found in the HZ. Scalo et al. (2007) provide a
effects of clouds (Kitzmann et al. 2013), plane- detailed review of habitability issues. The HZ is
tary orbit (e.g., Williams and Pollard 2003), stud- rather close to the star (typically ~0.2 AU) so that
ies of climate evolution (e.g., Selsis et al. 2007), Earth-like exoplanets could be tidally locked,
and (of particular interest) planetary mass (see slow rotators. The 3D modeling study of Joshi
“Super-Earths” below) and ocean mass (Abe (2003) suggested that atmospheres of such
et al. 2011) (see “Waterworlds and Desertworlds” worlds could effectively transport heat from the
below). dayside to the nightside, hence maintaining hab-
itable conditions. Joshi and Haberle (2012)
Super-Earths suggested that including the spectral dependence
These objects (with mass M such that of the snow-ice albedo for modeled planets
MEarth<M<10 MEarth) are already starting to be orbiting red dwarf stars led to an increase in the
found in the HZ (see below). There exists a lively outer HZ by up to 30 % due to a weaker snow-ice
discussion as to their ability to maintain plate albedo feedback. Shields et al. (2013) investi-
tectonics (e.g., Noack and Breuer 2013) which gated similar effects, finding that Earth-like
would favor habitability and affects atmospheric planets in the HZ of M stars could withstand a
mass and composition via outgassing. The reduction of up to ~19 % in incoming insolation
enhanced planetary mass of super-Earths favors without “going snowball” – partly because of the
slower thermal escape of the original (proto) weaker ice-albedo feedback. Kite et al. (2011)
H2-rich atmosphere. For example, Pierrehumbert modeled potentially destabilizing (negative)
206 Atmospheric Habitability
climate feedbacks involving weathering and the importance of coupling climate and photo-
pressure on tidally locked planets. Decreasing chemistry when calculating biosignature abun-
dayside pressure led to a warming (air is less dances. For example, they showed that key
efficiently transported to the nightside) which in bioindicators such as O3 and important green-
turn led to a faster weathering rate, which further house gases such as CH4 respond sensitively to
decreases the atmospheric pressure. Leconte the stellar input spectrum and the planet’s posi-
et al. (2013) studied (near) tidally locked planets tion in the HZ. The 3D study of Godolt (2012)
close to the inner HZ boundary and discussed two discussed a climate feedback which could warm
opposing 3D effects – firstly, water evaporated on (cool) the atmospheres of Earth-like planets
the dayside (suggesting a runaway climate effect) orbiting in the HZ of K stars (F stars) – they
but secondly, water froze out on the nightside found that enhanced (suppressed) incoming
(opposing the runaway effect). Yang longwave infrared radiation for the M dwarf star
et al. (2013) analyzed a cloud feedback mecha- scenario favors stronger (weaker) planetary
nism in their 3D model study which increased the atmospheric heating, and the effect is strength-
planetary albedo and hence expanded the inner ened by enhanced (decreased) water vapor evap-
HZ boundary toward the host star for Earth-like oration and a stimulated (suppressed)
planets orbiting M dwarf stars. A recent highlight hydrological cycle. Additional exoplanets
is, e.g., planetary studies of the GJ-667 system orbiting K stars which could be habitable are
(e.g., Anglada-Escudé et al. 2013). Finally, the recently emerging, e.g., HD85512b (Pepe
recent Kepler results are starting to suggest that et al. 2011).
planets in the HZ could be quite common.
Although Earth-like planets in the HZ of Habitability Beyond the Classical HZ
M dwarfs are in some ways favored objects, The main motivation here is to estimate how
there are nevertheless some aspects which could common is habitability (in all its possible
oppose the habitability of rocky planets orbiting forms) in the Universe. Most studies to date
in the HZ of M dwarf stars. Lammer et al. (2007), have focused on the Earth (“what we know”).
for example, suggested efficient escape of even On the other hand, one should clearly keep an
hundreds of bars of CO2 or/and N2 atmospheres open mind on the new and exciting results to
due to nonthermal escape processes for planets in come. Beyond the classical definitions, one
the close-in HZ of M dwarf stars especially dur- might include alternatives to the classical requi-
ing extended early, active stellar phases – this sites of life, including energy coming, e.g., not
aspect needs further investigation. Also, possibly only from main sequence stars or from white
weak (or even absent) magnetospheres for Earth- dwarf HZs which move continuously inward
like planets in the HZ of M dwarf stars could with time. Also, Neubauer et al. (2011) calculated
result in strong bombardment of the planetary the (much-expanded) HZ based on non-water
atmosphere by stellar and galactic cosmic rays. solvents.
Associated climate and photochemical effects
have been discussed by, e.g., Grenfell
et al. (2012, 2013) and Segura et al. (2010). Applications
Atmospheric Habitability of K and F Stars Key applications of the above theoretical studies
K stars are also favored targets for exoplanet (e.g., in an exoplanet context) are, e.g., to gener-
search missions, being rather cool, but unlike ate scientific debate by helping to understand the
the case of the M dwarfs, planets in their HZs main physical responses (e.g., between atmo-
are unlikely to be tidally locked. Segura spheric dynamics, climate and composition, and
et al. (2003) and Grenfell et al. (2007) modeled their effect on habitability), to produce scientific
the climate and photochemistry of Earth-like databases (e.g., theoretical atmospheric spectral
planets in the HZ of K and F stars and showed catalogs) which help interpret current and
future-planned data, and to drive future instru- cosmic rays for Earth-like planets in the habitable
mentation design by predicting the range of zone of M dwarf stars. Astrobiology
12(12):1109–1122
observed signals expected and the associated Grenfell JL, Gebauer S, Godolt M, Palczynski K, Rauer A
signal-to-noise ratio. H (2013) Potential biosignatures in Super-Earth-
atmospheres, II. Photochemical responses. Astrobiol-
ogy 13(5):415–438
Hart MH (1978) The evolution of the atmosphere of the
Future Directions Earth. Icarus 33:23–39
Huang SS (1960) Life outside the solar system. Sci Am
Theoretical studies to assess our understanding of 202:55–63
habitability are needed now, to prepare the way Joshi M (2003) Climate model studies of synchronously
rotating planets. Astrobiology 3(2):415–427
for next-generation missions such as the James Joshi MM, Haberle RM (2012) Suppression of the water
Webb Space Telescope (JWST) and the Euro- ice and snow albedo feedback on planets orbiting red
pean Extremely Large Telescope (E-ELT) dwarf stars and subsequent widening of the habitable
which could detect the first exoplanetary atmo- zone. Astrobiology 12(1):3–8
Kasting JF, Toon OB, Pollack JB (1988) How climate
spheric biosignatures (although this task is very evolved on the terrestrial planets. Sci Am 256:90–97
challenging). Kasting JF, Whitmire DP, Reynolds RT (1993) Habitable
zones around main sequence stars. Icarus 101:108
Kite E, Gaidos E, Manga M (2011) Climate instability on
tidally locked exoplanets. Astrophys J 743(1):1–12
See Also Kitzmann D, Patzer ABC, von Paris P, Godolt M,
Stracke B, Gebauer S, Grenfell JL, Rauer H (2010)
▶ Habitable Zone Clouds in the atmospheres of extrasolar planets.
▶ Paleosols I. Climatic effects of multi-layered clouds for Earth-
like planets and implications for habitable zones.
▶ Plate Tectonics Astron Astrophys 511:A66
Kitzmann D, Patzer B, Rauer H (2013) Clouds in the
atmospheres of extrasolar planets. IV. On the scatter-
References and Further Reading ing greenhouse effect of CO2 ice particles: numerical
radiative transfer studies. A&A 557:A6
Abe A, Abe-Ouchi A, Sleep NH, Zahnle KJ (2011) Habit- Kopparapu R, Ramirez R, Kasting JF, Eymet, Robinson T,
able zone limits for dry planets. Astrobiology Mahadevan S, Ryan C, Terrien RC, Domagal-
11(5):443–460 Goldman S, Meadows V, Deshpande R (2013) Habit-
Anglada-Escudé G, Tuomi M, Gerlach E, Barnes R, able zones around main-sequence stars: new estimates.
Heller R et al (2013) A&A 556:126 Astrophys J 770:82
Dole SH (1964) Habitable planets for man, 1st edn. Lammer H, Lichtenegger HIM, Kulikov Y, Griessmeier
Blaisdell, New York. ISBN 0-444-00092-5 JM, Terada N, et al (2007) Coronal mass ejection
Godolt M (2012) 3D climate modeling of Earth-like extra- (CME) activity of low mass stars as an important factor
solar plants orbiting different types of central star, PhD for the habitability of terrestrial exoplanets, 7(1),
thesis, Technische Universität Berlin (TUB), Germany 167–184
Goldblatt C, Matthews AJ, Claire MW, Lenton TM, Wat- Leconte J, Forget F, Charney B et al (2013) 3D climate
son AJ, Zahnle KJ (2009) Nitrogen enhanced green- modeling of close-in land planets: circulation patterns,
house warming on early Earth. Nat Geosci 2:891–896 climate moist bistability and habitability. A&A 554:
Grenfell JL, Stracke B, von Paris P, Patzer ABC, Titz R, A69
Segura A, Rauer H (2007) The response of atmo- Neubauer D, Vrtala A, Leitner JJ, Ferneis MG,
spheric chemistry on Earth-like planets around F, Hitzenberger R (2011) Development of a model to
G and K stars to small variations in orbital distance. compute the extension of life supporting zones for
Planet Space Sci 55:661–671 Earth-like exoplanets. Orig Life Evol Biosph
Grenfell JL, Gebauer S, von Paris P, Godolt M, Hedelt P, 41(6):545–552
Patzer ABC, Stracke B, Rauer H (2011) Sensitivity of Noack L, Breuer D (2013) First- and second-order Frank-
biomarkers to changes in chemical emissions in Kamenetskii approximation applied to temperature-,
Earth’s proterozoic atmosphere. Icarus 211:81–88 pressure- and stress-dependent rheology. Geophys
Grenfell JL, Griessmeier J-M, von Paris P, Patzer ABC, J Int 195:27–46
Lammer H, Stracke B, Gebauer S, Schreier F, Rauer Pepe F, Lovis C, Segransan D, Benz W, Bouchy F,
H (2012) Response of atmospheric biomarkers to Dumusque X, Mayor M, Queloz D, Santons NC,
NOx-induced photochemistry generated by stellar Udry S (2011) The HARPS search for Earth-like
208 Atmospheric Particles
planets in the habitable zone. I. Very low-mass planets

around HD20794, HD85512, and HD192310. A&A Atmospheric Redox Change
534:A58
Pierrehumbert R, Gaidos S (2011) Hydrogen greenhouse
planets beyond the habitable zone. Astrophys J 734: ▶ Oxygenation of the Earth’s Atmosphere
L13
Scalo J, Kaltenegger L, Segura A, Fridlund M, Ribas
I et al (2007) A re-appraisal of the habitability of
planets around M-dwarf stars. Astrobiology 7:30–65 Atomic Fine Structure Cooling
Segura A, Krelove K, Kasting JF, Sommerlatt D,
Meadows V, Crisp D, Cohen M, Mlawer E (2003) Steven B. Charnley
Ozone concentrations and ultraviolet fluxes on Earth- Solar System Exploration Division, Code 691,
like planets around other stars. Astrobiology
3:689–708 Astrochemistry Laboratory, NASA Goddard
Segura A, Walkowicz LM, Meadows V, Kasting J, Space Flight Center, Greenbelt, MD, USA
Hawley S (2010) The effect of a strong stellar flare
on the atmospheric chemistry of an Earth-like planet
orbiting an M-dwarf. Astrobiology 10(7):751–771.
doi:10.1089/ast.2009.0376
Definition
Selsis F, Kasting JF, Levrard B, Paillet J, Ribas I, Delfosse
X (2007) Habitable planets around the star Gliese 581? Atomic fine structure cooling is one of the processes
A&A 476:1373–1387 by which ▶ interstellar medium material can lose
Shields AL, Bitz CM, Meadows VS, Joshi MM, Robinson
TD (2013) The effect of host star spectral energy
thermal energy. Atomic and molecular hydrogen
distribution and ice-albedo feedback on the climate collisions excite the fine structure components of
of extrasolar planets. Astrobiology 13(8):715–739 ground state terms in atoms and atomic ions – e.g.,
Som SM, Catling DC, Harnmeijer JP, Polivka JP, Buick in neutral carbon and oxygen atoms and in C+ – and
R (2012) Air density 2.7 billion years ago limited to
less than twice modern levels by fossil raindrop
these subsequently decay radiatively with the
imprints. Nature. doi:10.1038/nature10890 escaping photon leading to a net cooling of the gas.
Steele A, Beaty DW, Amend J, Anderson R, Beegle L,
Benning L et al (2005) The astrobiology field labora-
tory, MEPAG white paper, 75 pp Dec 2005 See Also
von Paris P, Gebauer S, Godolt M, Grenfell JL, Hedelt P,
Kitzmann D, Patzer ABC, Rauer H, Stracke B (2010)
The extrasolar planet GL 581d: a potentially habitable ▶ Interstellar Medium
planet? A&A 522:A23 ▶ Molecular Line Cooling
von Paris P, Grenfell JL, Rauer H, Stock J (2013) ▶ Photodissociation Region
N2-associated warming on early Mars. Planet Space
Sci 82:149–154
Williams DM, Pollard D (2003) Extraordinary climates of
earth-like planets: three-dimensional climate simula-
tions at extreme obliquity. Int J Astrobiol 2:1–19 Atomic Nitrogen
Woolley R, Epps EA, Penston MJ, Pocock SB (1970) Roy
Obs Ann 5:227
Wordsworth R, Pierrehumbert RT (2013) Water loss from ▶ Nitrogen
terrestrial planets with CO2-rich atmospheres.
Astrophys J 778:154
Yang J, Cowan NB, Abbot DS (2013) Stabilizing cloud
feedback dramatically expands the habitable zone of ATP
tidally locked planets. Astrophys J Lett 771(2):L45
Shin Miyakawa
Atmospheric Particles Synonyms
▶ Aerosols Adenosine triphosphate; pppA

ATPase 209
Definition ▶ electrochemical potential of an ion (H+ or Na+)

into chemical energy of the system ATP-ADP.
ATP is one of the four activated nucleotides ATP synthases have a universal phylogenetic A
incorporated into ▶ RNA by RNA polymerases. distribution, and hence, it is supposed that
It is also extremely important in biochemistry as a they were present in the universal common
high-energy molecule that releases the energy ancestor.
needed for many metabolic reactions by coupling
hydrolysis into ADP and inorganic phosphate, or
AMP and pyrophosphate, with various synthetic
and mechanical processes. The formation of ATP See Also
in living organisms takes place through two main
biochemical pathways, substrate-level phosphor- ▶ ATP
ylation (in which phosphate is transferred to ADP ▶ Bioenergetics
from activated phosphorylated intermediates) ▶ Electrochemical Potential
and membrane-associated processes (found in ▶ Last Universal Common Ancestor
respiration and photosynthesis). Although prebi- ▶ Transduction
otic phosphorylation may have led to ATP from
adenosine, ATP is usually not considered to have
been abundant in prebiotic environments, and
polyphosphates have been proposed as alterna-
ATPase
tive primordial energy carriers.
José Pascual Abad
Facultad de Ciencias, Departamento de Biologı́a
See Also
Molecular, Universidad Autónoma de Madrid,
Cantoblanco, Madrid, Spain
▶ Adenine
▶ Nucleotide
▶ RNA
Keywords
A-ATPase; ABC-transporter; ATP synthase;

F-ATPase; Multidrug transporter; P-ATPase;
ATP Phosphohydrolase
Proton gradient; Proton motive force; Transmem-
brane ATPase; V-ATPase
▶ ATPase
Synonyms
ATP Synthase
Adenosine 50 -triphosphatase; Adenosine
Juli Peretó
triphosphatase; ATP phosphohydrolase
Evolutiva, Universitat de Valéncia, Valéncia,
Spain
Definition
Definition Enzymatic activity that catalyzes the decomposi-

tion of adenosine triphosphate (ATP) into aden-
ATP synthase is an enzymatic complex responsi- osine diphosphate (ADP) and phosphate (Pi).
ble for converting the energy of a transmembrane A protein with ATPase activity.
210 ATPase
Overview enzymes are homologues. In F-ATPases, a FO

complex of proteins integral to the membrane is
The term ATPase applies to the activity of any a transmembrane proton translocase, and the F1
enzyme’s ability to decompose ATP, including protein complex is a molecular machine that can
metabolic enzymes involved in anabolic pro- either decompose ATP promoting the passage of
cesses that need energy, as well as enzymes pro- proton through FO uphill the gradient or trans-
moting transport across ▶ membrane. In the form the energy liberated by the passage of pro-
former case, the enzyme decomposes ATP using ton, downhill the gradient of protons, through FO
the energy liberated to do work; for instance, into chemical energy, thus synthesizing ATP
enzymes such as DNA helicases, RecA protein, from ADP and Pi. Both complexes consist of
AAA proteins, or the muscle contraction protein assemblies of several subunits that together
myosin hydrolyze ATP, applying the liberated form a rotary motor (Fig. 1). Binding of protons
free energy directly to their functions. In other to FO causes the rotation of some of the subunits,
cases, i.e., glutamine synthetase, decomposition which together with the subunits of F1 conform
of ATP is performed in two steps, helping to three binding sites for ADP and Pi. The complex
perform chemical work, phosphorylating a sub- undergoes rotationally conformation changes
strate or enzyme in a first step, and releasing the through different affinity states, in one of which
Pi in a second step. Sometimes this type of reac- ADP and Pi can react, before the synthesized
tion involves the transfer of pyrophosphoryl or ATP is released (rotational catalysis).
adenylyl groups instead. There are many known
ATPases, which promote ATP hydrolysis
coupled to transport of solutes across membranes
into another compartment. These are integral β
α α
membrane proteins called transmembrane β β β
α ATP
ATPases, and there are many different types
found in all kind of organisms, performing func-
tions in which energetic exchanges are quantita-
tively quite relevant. There are at least four
general types of this kind of ATPases: F, V, ADP + Pi
ABC-multidrug transporters, and P. b2
F-ATPases, present in plasma membrane of
bacteria, mitochondrial inner membrane, and thy- H+ γ
lakoid membranes of chloroplasts of photosyn-
thetic eukaryotes, are reversible enzymes that are ε
Cytoplasm
able to promote the generation of a proton motive
force at the expense of ATP hydrolysis or to
synthesize ATP from ADP and Pi using a a
preexisting proton gradient. In this type of func- Membrane
tion, the enzyme is more appropriately named
▶ ATP synthase. This process of ATP synthesis C10−15
is often called oxidative phosphorylation. In the
case of fermenting bacteria which lack an elec-
tron transport chain (to produce a proton motive
force) and cannot perform oxidative phosphory-
H+
lation, the enzyme acts as a real ATPase produc-
F-ATPase (ATP synthase)
ing a proton gradient, which is used to energize
other transport processes. F-ATPases are struc- ATPase, Fig. 1 Model of the structure and function of
turally related to V-ATPases since both kinds of F-ATPases
ATPase 211
B
structurally and phylogenetically related to the
A A V-ATPases.
B B In some anaerobic bacteria, extremophiles
E A ATP A
(particularly thermophilic and alkaliphilic), and
Archaea, the coupling ion used by F and V-types
ATPases is Na+ instead or in addition to H+, and a
sodium motive force (SMF) substitutes the
ADP + Pi
G2 ▶ proton motive force, (PMF). Certain studies
suggest that the Na+-dependent ATPases came
H+ first in terms of evolutionary history; however,
D
F some others consider them as secondary adapta-
tion to survival in extreme environments.
Cytoplasm d/C The ATP-binding cassette (ABC) multidrug
transporters constitute a large and ubiquitous
superfamily of integral membrane proteins that
a/I are responsible for the ATP-powered transloca-
Membrane
tion of many substrates across membranes,
existing as both importers and exporters.
c/K6−13
Importers have, to this date, only been found in
Lumen
prokaryotes, whereas exporter-type ABC trans-
porters are expressed in the three domains of life.
The highly conserved ABC structural domains of
H+ ABC transporters provide the nucleotide-
V-ATPase dependent engine that drives transport. By con-
trast, the transmembrane domains that generate
ATPase, Fig. 2 Model of the structure and function of the translocation pathway are more variable. In
V-ATPases
importers, the ABC domains are in separate sub-
units from the transmembrane ones (TMD) while
in exporters are another domain of the transporter
V-ATPases, present in lysosomal or protein (Fig. 3). In prokaryotes, the importers
endosomal secretory vesicle membranes, are provide the cell with hydrophilic nutrients,
proton-transporting proteins that generate a while exporters remove different xenobiotics
lower pH inside the vesicles, thus activating the and hydrophobic substances from the cell. The
hydrolytic enzymes present in such compart- prokaryotic exporters are responsible for the
ments. V-ATPases are also found in the plasma efflux of drugs that cause certain antibiotic resis-
membranes of a wide variety of animal cells, tances, and the eukaryotic ones can prevent the
including certain tumor cells, where they are inhibitory effect of anticancer drugs by
involved in processes such as pH homeostasis or preventing their accumulation within the tumor
metastasis. V-ATPases have also been found in cells. For functioning, the importers require a
plants and fungi. The structures of these enzymes substrate-binding protein (SBP).
are more complex than those of P-ATPases and P-ATPases are enzymes of plasma or sarco-
include a transmembrane proteins complex (VO), plasmic reticulum that promote cation transport;
which works as a proton translocase, and a these enzymes are reversibly phosphorylated by
peripheral domain (V1), with the ATPase activ- ATP. All members of this family have certain
ity. The better known is that of yeast in which VO amino acid sequence conservation, particularly
seems to contain six and V1 eight different sub- the Asp residue that suffers the phosphorylation/
units (Fig. 2). The ATPases of Archaea dephosphorylation process, and are inhibited by
(sometimes referred to as A-ATPases) are vanadate. Some are a unique polypeptide with
212 ATPase
Exporter Importer
SBP
Hydrophilic nutrients
Membrane TMD TMD Membrane TMD TMD
Cytoplasm Cytoplasm
ABC ABC ABC ABC
ATP ADP + Pi ATP ADP + Pi
Hydrophobic substances
ABC-multidrug transporters
ATPase, Fig. 3 Model of the structure and function of ABC-multidrug exporters and importers. TMD transmembrane
domain, ABC ATP-binding cassette, SBP substrate-binding protein
multiple membrane-spanning regions, while

others have a second subunit. This type of
enzyme is widely distributed and includes repre-
sentatives in animals (Na+K+-ATPase, Ca2+-
ATPase, H+K+-ATPase), plants and fungi
(plasma membrane H+-ATPase), and bacteria
(Cd2+, Hg2+, or Cu2+ ATPases). Albeit their
diversity in ion specificity, all P-ATPases are
homologues (Fig. 4).
Basic Methodology
The main methodologies used for the studies on

ATPases include crystallization and structure
determination by X-ray diffraction; nuclear mag-
ATPase, Fig. 4 Model of the structure and function of
netic resonance (NMR) of particular subunits or the Ca+-ATPase, a P-ATPase
complexes; electron and fluorescence micros-
copy including visualization of the rotational
movement of F1-ATPase, after attaching the Key Research Findings
rotating subunit to a fluorescent actin filament;
and kinetic and binding studies of the enzyme Boyer’s group proposed in 1973 that the step
activities, as well as genetic studies based on the requiring energy in the synthesis of ATP by
effect of protein sequence modifications. ATP synthase was the release of the ATP
ATPase 213
molecule from the enzyme. Three years later, the Buch-Pedersen MJ, Pedersen BP, Veierskov B, Nissen P,
binding change mechanism was proposed by this Palmgren MG (2009) Protons and how they are
transported by proton pumps. Eur J Physiol
group. The first X-ray structure of an incomplete 457:573–579 A
F1 complex was published by Walker’s group in Gr€uber G, Marshansky V (2008) New insights into
1994. In 1997, Yoshida’s group was able to record structure-function relationships between archeal ATP
the movement of the F1 complex during ATP synthase (A1A0) and vacuolar type ATPase (V1V0).
Bioessays 30:1096–1109
hydrolysis, now including the gamma subunit Itoh H, Takahashi A, Adachi K, Noji H, Yasuda R,
(the rotating shaft of the enzyme), after attaching Yoshida M, Kinosita K Jr (2004) Mechanically driven
individual complexes to a surface and looking to a ATP synthesis by F1-ATPase. Nature 427:465–468
fluorescent actin filament attached to the g subunit. Jefferies KC, Cipriano DJ, Forgac M (2008) Function,
structure and regulation of the vacuolar (H+)-
In 2004, the mechanically driven synthesis of ATP ATPases. Arch Biochem Biophys 476:33–42
by the F1 complex was also demonstrated in exper- Kayalar C, Rosing J, Boyer PD (1977) An alternating site
iments in which the g subunit inserted in the F1 sequence for oxidative phosphorylation suggested by
complex was induced mechanically to rotate in the measurement of substrate binding patterns and
exchange reaction inhibitions. J Biol Chem
presence of ADP and phosphate and ATP was 252:2486–2491
afterward released to the medium. Krah A, Pogoryelov D, Meier T, Faraldo-Gómez JD
(2010) On the structure of the proton-binding site in
the Fo rotor of chloroplast ATP synthases. J Mol Biol
395:20–27
Future Directions K€uhlbrandt W (2004) Biology, structure and mechanism
of P-type ATPases. Nat Rev Mol Cell Biol 5:282–295
Although we begin to have a good view of how Locher KP (2009) Structure and mechanism of
ATPases work, deeper structural analyses are ATP-binding cassette transporters. Philos Trans
R Soc B 364:239–245
needed to determine the mechanisms of ions pass Madigan MT, Martinko JM, Dunlap PV, Clark DP
through the ATPases domains and of torque gener- (2008) Brock biology of microorganisms, 12th edn.
ation. Comparison of the structures of ATPases of Benjamin Cumming, San Francisco
different origins should also help in determining the Mulkidjanian AY, Makarova KS, Galperin MY, Koonin
EV (2007) Inventing the dynamo machine: the evolu-
most relevant aspects of this issue. In the tion of the F-type and V-type ATPases. Nat Rev
astrobiological field, the ions that could have been Microbiol 5:892–899
used by the first ATPases, in the origin of life, are Mulkidjanian AY, Galperin MY, Makarova KS, Wolf YI,
still controversial, and thus research in this field is of Koonin EV (2008) Evolutionary primacy of sodium
bioenergetics. Biol Direct 3:13
great interest and will have to be addressed shortly. Mulkidjanian AY, Galperin MY, Koonin EV (2009)
Co-evolution of primordial membranes and membrane
See Also proteins. Trends Biochem Sci 34:206–215
Nelson DL, Cox MM (2009) Lehninger principles of bio-
chemistry, 5th edn. WH Freeman, New York
▶ ATP Synthase Noji H, Yasuda R, Yoshida M, Kinosita K Jr (1997) Direct
▶ Bioenergetics observation of the rotation of F1-ATPase. Nature
▶ Membrane 386:299–302
▶ Proton Motive Force Pedersen PL (2007) Transport ATPases into the year
2008: a brief overview related to types, structures,
functions and roles in health and disease. J Bioenerg
Biomembr 39:349–355
References and Further Reading Pogoryelov D, Yildiz Ö, Faraldo-Gómez JD, Meier
T (2009) High-resolution structure of the rotor ring of
Abrahams JP, Leslie AG, Lutter R, Walker JE a proton-dependent ATP synthase. Nat Struct Mol Biol
(1994) Structure at 2.8 Å resolution of F1-ATPase 16:1068–1073
from bovine heart mitochondria. Nature 370:621–628 Rees DC, Johnson E, Lewinson O (2009) ABC trans-
Boyer PD, Cross RL, Momsen W (1973) A new concept porters: the power to change. Nat Rev Mol Cell Biol
for energy coupling in oxidative phosphorylation 10:218–227
based on a molecular explanation of the oxygen Saroussi S, Nelson N (2009) Vacuolar H + -ATPase an
exchange reactions. Proc Natl Acad Sci U S A enzyme for all seasons. Eur J Physiol 457:581–587
70:2837–2839
214 AU
Definition
AU
Autocatalysis is catalysis by one or more of the
Daniel Rouan products of a reaction. Autocatalysis is often seen
LESIA, Observatoire Paris-Site de Meudon, as the minimal requirement for the emergence of
Meudon, France ▶ life, as it is at the core of modern biogenetic
theories based on genetic replicators, metabolic
networks, and containment reproducers. Autoca-
Synonym talysis is one of the pathways for chiral symmetry
breaking and is also responsible for the formation
Astronomical unit of patterns and ordered periodic behavior in
chemical reactions. While autocatalytic phenom-
ena have been observed in fields as diverse as cell
Definition biology and nonlinear physics, their general
study is the subject of the emergent field of “Sys-
AU stands for astronomical unit. It is the average tems Chemistry.”
Earth-Sun distance, corresponding to
149.6 106 km (or approximately 8 light-min
or 100 Sun diameters). This is one of the basic Overview
units of astronomy. Other distance units (pc, kpc,
etc.) are derived from it. Autocatalytic reactions are described by the
equation:
See Also pP þ aA þ . . . ! ðp þ 1ÞP þ . . . (1)
▶ Parsec where a and p are reaction orders, P is the

autocatalyst, and A is a precursor molecule. The
autocatalytic reaction order p determines the
“explosivity” of the product growth, which
increases from parabolic (p = ½) to exponential
Austrian Space Agency, Austria (p = 1) and hyperbolic (p = 2). Each type of
growth has distinct evolutionary consequences.
▶ ASA
Competition between two or more
autocatalysts – for a common resource constitu-
ent A – leads to coexistence (“survival of every-
body”) for parabolic, selection (“survival of the
Autocatalysis fittest”) for exponential, and fixation (“survival of
the least common”) for hyperbolic growth.
Olga Taran1 and G€unter von Kiedrowski2 Typically, the concentration-time curve of for-
1
Chemistry and Chemical Biology, Harvard mation of an autocatalyst has a sigmoidal form
University, Cambridge, MA, USA (see Fig. 1), which is the result of a self-
2
Lehrstuhl f€
ur Organische Chemie I, Ruhr- accelerating growth phase followed by decelera-
Universität Bochum, Bochum, NRW, Germany tion due to the consumption of precursors.
Coupling of autocatalytic cycles with other
reactions including autocatalytic ones can lead
Keywords to oscillating behavior and spatiotemporal pattern
formation. One example is the well-known
Catalysis; Origin of life theories; Self-replication Belousov-Zhabotinsky reaction.
Autocatalysis 215
a 10 b
8
A
Concentration
6
0
0 5 10 15 20
Time
nA Ap
c
BA
BB
AA
AB
Bp nB
Autocatalysis, Fig. 1 (a) Typical sigmoidal concentration oscillations of ferroin (Fe (II)) and ferritin
concentration-time curve observed in many autocatalytic (Fe (III)) complexes in Belousov-Zhabotinsky reaction.
reactions. Initially, the reaction rate is slow, while the Reaction with limited diffusion creates spatial patterns as
catalyst concentration is low. The reaction rate increases observed in this photo. (c) Scheme of a simple four-
as more products are formed and then slows down when component cross-catalytic network. Precursors A and
the regents are consumed or inhibition by template takes B can form two autocatalysts AB and BA or a group of
place. The character of the growth phase can be parabolic, cross-catalysts AA and BB. The scheme represents the
exponential, or hyperbolic, according to the reaction reaction mechanism reported by Sievers and von
mechanism involved in the process. (b) Periodic Kiedrowski (1994)
Autocatalysis can be found in both catabolic based on a templating principle according to

reactions (e.g., hydrolysis of simple esters) and which the product transfers its blueprint as con-
anabolic reactions (e.g., template replicators). stitutional information. It has been demonstrated
Feedback may result from simple autocatalysis using nucleic acids, peptides, and small organic
by one product, cross-catalysis between two molecules as templates.
products, or an “autocatalytic set” of molecules Metabolic theories call for cycles that generate
forming a network of mutually catalytic larger molecules from small building blocks in an
interactions. anabolic direction, whereas the autocatalysis
Many genetic theories of the origin of life itself results from a catabolic split to yield
(e.g., the “▶ RNA world” model) demand self- medium-sized intermediates (as in the
replication of nucleic acids in the absence of “▶ Formose reaction”).
enzymes. Self-replication means autocatalysis Autopoietic theories require self-reproduction
plus information transfer. The latter is usually of a container as well as the included
216 Automaton, Chemical
components. Experimental examples have been

found in reactions where lipid molecules forming Automaton, Chemical
micelles or vesicles are generated from non-lipid
precursors via phase-transfer autocatalysis. André Brack
Several modern theories for the origin of (bio) Centre de Biophysique Moléculaire CNRS,
molecular homochirality are supported by exam- Orléans Cedex 2, France
ples of enantioselective autocatalysis and chiral
symmetry breaking.
Current research challenges in the field of Keywords
Systems Chemistry are programmed design of
autocatalytic systems and their integration into Algorithmic chemistry; Autopoiesis; Chemical
dynamic supersystems. reaction network; Chemoton; Minimal
metabolism
See Also
Synonym
▶ Autocatalysis
Self-replication
▶ Evolution, Molecular
▶ Formose Reaction
▶ Hypercycle
Definition
▶ Life
▶ RNA World
There is a remarkable gap between complex
▶ Self-Replication
chemical systems that generate self-organizing
▶ Template-Directed Polymerization
patterns (e.g., oscillations in Belousov-
Zhabotinsky types of reactions) and the chemical
References and Further Reading self-construction that any living being achieves
through its metabolism. Chemical self-
Bachmann PA, Luisi PL, Lang J (1992) Autocatalytic self- organization mechanisms operate at many differ-
replicating micelles as models for prebiotic structures.
Nature 357:57–59
ent levels in biological systems, but the network
Blackmond D (2004) Asymmetric autocatalysis and its of continuous transformation processes underly-
implications for the origin of homochirality. Proc ing the constitution of an organism is something
Natl Acad Sci 101:5732–5736 clearly distinct. The idea of a chemical automa-
Eigen M, Schuster P (1979) The hypercycle – a principle
of natural self-organization. Springer, Berlin
ton addresses precisely this issue, trying to deter-
Epstein IR, Pojman JA (1998) An introduction to mine what an autocatalytic reaction network
nonlinear chemical dynamics: oscillations, waves, pat- requires to become an autonomous, self-
terns, and chaos. Oxford University Press, New York producing system. So far we lack experimental
See the Memorandum of Understanding of the Systems
Chemistry at European Cooperation in the Field of
evidence of chemical automata that are not living
Scientific and Technical Research (COST) site http:// metabolic systems, but synthetic biology and cur-
w3.cost.esf.org/index.php?id=189&action_number= rent projects to fabricate artificial cells may soon
CM0703 provide interesting insights to this problem,
Sievers D, von Kiedrowski G (1994) Self-replication of
complementary nucleotide-based oligomers. Nature
which has been extensively explored theoreti-
369:221–224 cally in the past.
Szathmary E (2006) The origin of replicators and repro-
ducers. Philos Trans R Soc Lond B Biol Sci
361:1761–1776
Vidonne A, Philp D (2009) Making molecules make
See Also
themselves – the chemistry of artificial replicators.
Eur J Org Chem 5:593–610 ▶ Autocatalysis
Autotroph 217
▶ Chemical Reaction Network Definition

▶ Self-Assembly
▶ Self-Replication Autopoiesis is a recurrent set of component pro- A
duction processes that creates a physical/topolog-
ical boundary, within which that set of processes
is continuously realized.
Benner SA, Ricardo A, Carrigan MA (2004) Is there a
common chemical model for life in the universe? Curr
Opin Chem Biol 8(6):672–689 History
Brack A, Troublé M (2010) Defining life: connecting robot-
ics and chemistry. Orig Life Evol Biosph 40:31–136
Bro P (1997) Chemical reaction automata. Complexity The term was originally proposed by
2(3):38–44 H. Maturana and F. Varela in the early 1970s
Eigen M, Schuster P (1979) The hypercycle: a principle of (from Greek: auto, “self,” and poiesis, “produc-
natural self-organization. Springer, New York tion”) and proposed as an abstract definition of
Fontana W (1992) Algorithmic chemistry. In: Langton
CG, Taylor C, Farmer JD, Rasmussen S (eds) ▶ life. According to these authors, an autopoietic
Artificial life II. Addison-Wesley, Redwood City, system is “organized as a network of processes of
pp 159–209 production (transformation and destruction) of
Ganti T (1975) Organization of chemical reactions into components which: (1) through their interactions
dividing and metabolizing units: the chemotons.
Biosystems 7:15–21 and transformations continuously regenerate
Kauffman S (1986) Autocatalytic sets of proteins. J Theor and realize the network of processes (relations)
Biol 119:1–24 that produce them; and (2) constitute it (the
Rosen R (1991) Life itself: a comprehensive inquiry into machine) as a concrete unity in the space in
the nature, origin and fabrication of life. Columbia
University Press, New York which they (the components) exist, by specifying
Ruiz-Mirazo K, Moreno A (2004) Basic autonomy as a the topological domain of its realization as such a
fundamental step in the synthesis of life. Artif Life network.”
10(3):235–259
Varela FJ, Maturana H, Uribe R (1974) Autopoiesis: the
organization of living systems, its characterization and
a model. Biosystems 5:187–196
See Also
▶ Automaton, Chemical
▶ Cell, Minimal
▶ Life
Autopoiesis ▶ Metabolism
Alvaro Moreno1 and Kepa Ruiz-Mirazo2

1
Departamento de Lógica y Filosofı́a de la
Ciencia, Universidad del Paı́s Vasco,
Autotroph
San Sebastián, Spain
2
Department of Logic and Philosophy of Science,
Elena González-Toril
FICE, UPV-EHU, Biophysics Research Unit
Laboratorio de Extremófilos, Centro de
(CSIC – UPV/EHU), Donostia, San Sebastián,
Astrobiologı́a (INTA-CSIC), Torrejón de Ardoz,
Spain
Madrid, Spain
Synonyms
Synonyms
Metabolic organization; Minimal autonomy;
Self-producing network Primary producer
218 Autotrophy
Definition Synonyms
Autotroph is an organism capable of Primary production

biosynthesizing all cell material from carbon
dioxide as the only carbon source. With respect
to energy, autotrophs can obtain it from two Definition
sources: (1) photoautotrophs from radiation
(sunlight) and (2) chemolithoautotrophs from Autotrophy is a lifestyle in which inorganic com-
the oxidation of reduced inorganic substrates. pounds provide for all nutritional needs of an
Autotrophs are capable of growth exclusively at organism. Implicit in this definition is the capac-
the expense of inorganic nutrients, and they are of ity of an organism to derive all cell carbon
vital importance in the cycling of inorganic com- from carbon dioxide. Energy can be derived
pounds on Earth including methanogens, which from two sources: (1) Photoautotrophs are pho-
produce methane from H2 and CO2, and nitrifiers, tosynthetic and obtain energy from sunlight.
which convert ammonia to nitrate. Autotrophs (2) Chemolithoautotrophs obtain energy by the
are the source of reduced carbon substrates for oxidation of inorganic substances. Table 1 shows
the heterotrophs. Autotrophs are key elements of a general classification of organisms on the basis
the ▶ carbon cycle. For this reason, autotrophic of the carbon and energy sources.
organisms are also called primary producers.
See Also Overview
▶ Autotrophy Autotrophs are capable of growth exclusively at

▶ Calvin-Benson Cycle the expense of inorganic nutrients, and they are
▶ Carbon Cycle, Biological vital in the cycling of inorganic compounds
▶ Carbon Dioxide (Alberts et al. 1994; Campbell and Reece 2002).
▶ Carbon Source Such autotrophs not only completely satisfied their
▶ Chemolithoautotroph own needs for reduced carbon monomers from
▶ Photoautotroph inorganic matter but could also feed the already
existing heterotrophs. Thus, autotrophic organisms
are also called primary producers. Carbon dioxide
that is fixed into organic compounds as a result of
Autotrophy autotrophic activity is available for consumption or
respiration by animals or heterotrophic microor-
Elena González-Toril1 and Juli Peretó2 ganism. The end products of respiration in hetero-
1
Laboratorio de Extremófilos, Centro de trophic organism are carbon dioxide, and this way
Astrobiologı́a (INTA-CSIC), Torrejón de Ardoz, the carbon cycle is completed (Alberts et al. 1994;
Madrid, Spain Campbell and Reece 2002). Now it is accepted that
2
Institut Cavanilles de Biodiversitat i Biologia autotrophy is an extremely important process on
Evolutiva, Universitat de València, València, Earth and autotrophic microorganisms, as primary
Spain producers, support the growth of non-autotrophic
organisms (Maier et al. 2000).
Keywords Photoautotrophs
A large number of microorganisms, as well as the
Calvin-Benson cycle; Carboxysomes; green plants, algae, and protists, are
Chemolithoautotrophy; CO2 fixation; phototrophic. They use light as energy source in
Photoautotrophy; Rubisco the process called photosynthesis. The result of
Autotrophy 219
Autotrophy, Table 1 Classification of metabolisms according to energy, reducing power, and carbon sources
Metabolism Energy source Reducing power Carbon source
Chemo-litho-autotrophic Oxidation of inorganic compounds Inorganic compounds CO2 A
Photo-litho-autotrophic Visible light Inorganic compounds CO2
Photo-organo-heterotrophic Visible light Organic compounds Organic compounds
Chemo-organo-heterotrophic Oxidation of organic compounds Organic compounds Organic compounds
this mechanism is the generation of a proton The pathways differ in the participating enzymes,
motive force that can be used in the synthesis of ATP, and reducing power requirements and car-
ATP and the synthesis of reducing power (e.g., bon isotope fractionation (Maier et al. 2000;
NADPH). Most phototrophs use energy conserved Madigan et al. 2003; Berg et al. 2010).
in ATP and electrons in NADPH for the assimila-
tion of carbon dioxide as the carbon source for 1. The Calvin-Benson cycle, discovered in the
biosynthesis. These phototrophs are called photo- 1950s in Melvin Calvin’s lab, starts with the
autotrophs. There are also phototrophs able to use condensation of a 5-C sugar (ribulose
organic compounds as carbon sources with light as 1,5-bisphosphate) with CO2 to yield two mol-
energy source; they are called photoheterotrophs ecules of 3-C (3-phosphoglycerate) (Fig. 1a).
(Table 1) (Campbell and Reece 2002; Maier From these molecules both the initial 5-C
et al. 2000; Madigan et al. 2003). sugar is regenerated and organic materials
are biosynthesized. The cycle is operative in
Chemolithoautotrophs plastids of plants, algae, and protists, as well
In the 1880s, Sergei Winogradsky (1856–1953) pro- as in cyanobacteria, some aerobic or faculta-
posed the concept of chemolithotrophy, the oxidative anaerobic proteobacteria, CO-oxidizing
tion of inorganic compounds as a source of energy mycobacteria, some iron- and sulfur-oxidizing
and electrons for the autotrophic growth. Studying firmicutes, and green sulfur bacteria. The abil-
sulfur bacteria (Beggiatoa and Thiothrix), he con- ity to fix carbon by this pathway is conferred
cluded that these organisms obtained their carbon by the activity of two enzymes (together with
from CO2 in air, and they were called autotrophs. fragments of central metabolism like gluco-
The discovery of autotrophy in chemolithotrophic neogenesis and the pentose phosphate path-
bacteria was of major significance in the advance of way): ribulose 1,5-bisphosphate carboxylase-
our understanding of cells physiology because it oxygenase (▶ Rubisco) and phosphoribu-
showed that CO2 could be converted to organic lokinase (Campbell and Reece 2002; Maier
carbon without photosynthesis. Chemolithotrophy et al. 2000; Madigan et al. 2003). Although
is shown by members of both the Bacteria and Rubisco activity has been detected in some
Archaea domains. Chemolithoautotrophs are impor- Archaea, it has not been possible to demon-
tant in the cycling of inorganic compounds in Earth, strate Calvin-Benson-dependent autotrophic
including methanogens, which produce methane, growth (Berg et al. 2010).
and nitrifiers, which convert ammonia to nitrate Several autotrophic prokaryotes that use
(Madigan et al. 2003; Ehrlich 2002). the Calvin-Benson cycle for CO2 fixation pro-
duce polyhedral cell inclusions called
carboxysomes. Carboxysomes are made of
Basic Methodology and Key Research polyhedral protein shells about 80–120 nm in
Findings diameter (Fig. 2). These compartments are
surrounded by a thin membrane and consist
Autotrophic Pathways of a tightly packed crystalline array of mole-
Six biochemical mechanisms are known for the cules of Rubisco (Tsai et al. 2007). Thus, the
autotrophic fixation of CO2 into cell material. carboxysomes would be a mechanism to
220 Autotrophy
Autotrophy, Fig. 1 The

diversity of autotrophic
pathways. A scheme of
stoichiometric
relationships in (a) the
Calvin-Benson cycle, (b)
the Arnon cycle, (c) the
Wood-Ljungdahl pathway,
(d) the hydroxypropionate
bicycle, and (e) the
hydroxypropionate/
dicarboxylate-
hydroxybutyrate cycles. In
each case, the identity of
the net product of
C fixation, the starting point
of biosynthesis, is indicated
Autotrophy, Fig. 2 (a) A thin-section electron micro- H. neapolitanus. The features visualized arise from the
graph of Halothiobacillus neapolitanus cells with distribution of stain around proteins forming the shell as
carboxysomes inside. In one of the cells shown, arrows well as around the Rubisco molecules that fill the
highlight the visible carboxysomes. (b) A negatively carboxysome interior. Scale bars indicate 100 nm
stained image of intact carboxysomes isolated from (Figure from Tsai et al. (2007))
increase the amount of Rubisco in the cell to prochlorophytes. They are not present in fac-
allow for higher rates of CO2 fixation. ultative autotrophs like purple anoxygenic
Carboxysomes have been found in obligately phototrophs, despite the fact that when these
chemolithotrophic sulfur-oxidizing bacteria, organisms grow as photoautotrophs, they use
nitrifying bacteria, cyanobacteria, and the Calvin-Benson cycle to fix CO2. Thus, the
Autotrophy 221
carboxysome may be an evolutionary adapta- 5. The hydroxypropionate-hydroxybutyrate

tion to life under strictly autotrophic condi- cycle occurs in some aerobic archaea, like
tions (Madigan et al. 2003). Sulfolobus. Albeit this pathway is formally A
2. In the green sulfur bacterium Chlorobium, the same as the 3-hydroxypropionate bicycle,
CO2 fixation occurs by a reversal the nonhomologous participating enzymes
(or reductive) citric acid cycle also known as indicate that both pathways evolved indepen-
Arnon cycle (Madigan et al. 2003; Evans dently in a remarkable case of evolutionary
et al. 1966). This is the analogue of a Krebs convergence in metabolism (Berg
cycle operating in reverse mode (Fig. 1b). et al. 2010) (Fig. 1e).
Since the Arnon cycle involves some enzymes 6. The dicarboxylate-hydroxybutyrate cycle
(e.g., the carboxylating and reducing steps) occurs in some anaerobic archaea like the
that are inhibited by oxygen, this pathway is Thermoproteales and Desulfurococcales. The
only found in microorganisms growing under pathway can be divided into two parts
anaerobic conditions. These include some (Fig. 1e): (1) one acetyl CoA, one CO2, and
proteobacteria, green sulfur bacteria, and one bicarbonate are converted into succinyl
microaerophilic bacteria like Aquifex (Maier CoA; (2) this C4 molecule is transformed
et al. 2000; Madigan et al. 2003). into two molecules of acetyl CoA (one serves
3. In some Gram-positive bacteria and as biosynthetic precursor, the other as acceptor
methanogenic archaea, the ability to synthe- of the cycle) (Berg et al. 2010).
size acetyl CoA from CO and/or CO2, the
Wood-Ljungdahl pathway, was identified
(Madigan et al. 2003; Ljundahl et al. 1965). Applications
One CO2 molecule is reduced to CO and
another one to a methyl group (attached to a Ecology
cofactor). Then, acetyl CoA is synthesized Autotrophs are present in all ecosystems. They
from CO and the methyl group (Fig. 1c). The take energy from the environment in the form of
key enzymes of this pathway are inhibited by sunlight or inorganic chemicals and use it to
oxygen; thus it is restricted to obligate anaer- create energy-rich molecules such as carbohy-
obic microorganisms. These include some drates. Thus, they meet their requirements easily
proteobacteria, planctomycetes, spirochaetes, and can be constitutive around the world. More-
and archaea (Maier et al. 2000; Madigan over, autotrophic organisms are primary pro-
et al. 2003; Berg et al. 2010). ducers and, as a consequence, they are at the
4. The 3-hydroxypropionate bicycle is present in pyramidal base of the ecosystems. Thus, hetero-
some green non-sulfur phototrophic bacteria trophs depend on autotrophs for the energy and
like Chloroflexus (Herter et al. 2002). raw materials they need (Maier et al. 2000). On
A succinyl CoA molecule is synthesized the other hand, autotrophs, as a consequence of
from acetyl CoA and two bicarbonate mole- their poor requirements, have greater adaptabil-
cules (Fig. 1d). Although the same intermedi- ity; thus, they are especially important in oligo-
ates as the hydroxypropionate- trophic environments, like oligotrophic lakes,
hydroxybutyrate cycle (see below) are used, glaciers and ice, acid waters, geothermal
most of the participating enzymes are differ- systems, etc.
ent. The final product of the cycle is Rio Tinto (Huelva, Southwestern Spain) is an
glyoxylate. Its assimilation requires a second example of an environment dominated by auto-
metabolic cycle. This pathway is restricted to trophic bacteria. This ecosystem is of great inter-
the family Chloroflexaceae and might repre- est for astrobiology (Fig. 3). It is an extreme
sent an early attempt of autotrophy in environment with a rather constant acidic pH
anoxygenic phototrophs (Maier et al. 2000; along the entire river and a high concentration
Madigan et al. 2003). of heavy metals. The extreme conditions of the
222 Autotrophy
Autotrophy, Fig. 3 Rio

Tinto as example of an
ecosystem dominated by
autotrophic
microorganisms
Tinto ecosystem are generated by the metabolic Whether the first autotrophs were chemosyn-
activity of chemolithotrophic microorganisms thetic or photosynthetic is currently a matter of
thriving in the rich complex sulfides of the Ibe- debate. One school of thought favors chemosyn-
rian Pyrite Belt. In this system, more than 70 % of thetic autotrophs in the form of methanogens,
the cells are affiliated to autotrophic bacteria which formed methane. Microorganisms
(iron-oxidizing bacteria), with only a minor frac- (methanogenic archaea) with such metabolism
tion corresponding to heterotrophic. The special exist today, and they are strict anaerobes. The
interest shows also autotrophic microalgae, pre- other school of thought favors photosynthetic
sent in the river, primary producer, together with prokaryotes in the bacterial domain as the first
iron-oxidizing bacteria, of the system (González- autotrophs. This notion is supported by the exis-
Toril et al. 2003). tence of the Warrawoona stromatolites, which is
around 3.5 billion years old. Those microfossils
Autotrophy and Early Evolution of Life have been interpreted, on the basis of comparison
The autotrophic metabolism has emerged inde- with modern counterparts, to have been formed
pendently several times during evolution, that is, by cyanobacteria. However, modern
it is a polyphyletic trait (Berg et al. 2010; Pereto cyanobacteria are aerobes. Because the primor-
et al. 1999). The Calvin-Benson cycle seems dial atmosphere at this time is thought to have
idiosyncratic to bacteria, whereas the Arnon been almost free of oxygen, the emergence of
cycle and the Wood-Ljungdahl pathway show a anaerobic photosynthetic bacteria, of which mod-
wider phylogenetic distribution. The different ern purple and green bacteria must be a counter-
versions of the hydroxypropionate pathway part, must have preceded that of cyanobacteria
likely emerged independently (Berg et al. 2010). (Campbell and Reece 2002).
At this moment, phylogenetic analysis of the On the other hand, in 1988, Wächtershäuser
participating enzymes does not allow us to infer proposed the surface metabolism theory for the
which one is the older pathway. origin of life (Wächtershäuser 1988). According
Mainstream hypothesis on the ▶ origin of life to it, life arose as a form of autocatalytic
postulates that the first prokaryotes were anaero- two-dimensional chemolithotrophic metabolism
bic heterotrophs, for example, fermenters. In the on a pyrite surface, using the energy and electrons
beginning, they may have fed on externally avail- of the anaerobic synthesis of FeS2 (pyrite) from
able abiotic organic molecules, either synthe- FeS and H2S. According to this proposal, the
sized on Earth or delivered by extraterrestrial ancestral carbon fixation pathway would be a
bodies. It is generally supposed that autotrophic primitive version of the Arnon cycle
metabolism emerged latter. (Wächtershäuser 1990). The Wood-Ljungdahl
Azulmin 223
pathway has also been proposed as a candidate of Pereto J, Velasco AM, Becerra A, Lazcano A (1999)
the older autotrophic mechanism (Pereto Comparative biochemistry of CO2 fixation and the
evolution of autotrophy. Int Microbiol 2:3–10
et al. 1999; Russell and Martin 2004). The lack Russell MJ, Martin W (2004) The rocky roots of the A
of experimental evidences is the weaker aspect of acetyl-CoA pathway. Trends Biochem Sci 29:358–363
the autotrophic hypothesis on the origin of life. Tsai Y, Sawaya MR, Cannon GC, Cai F, Williams EB,
Heinhorst S, Kerfeld CA, Yeates TO (2007) Structural
analysis of CsoS1A and the protein shell of the
Halothiobacillus neapolitanus carboxysome. PLoS
See Also Biol 5(6):e144
Wächtershäuser G (1988) Before enzymes and templates:
▶ Calvin-Benson Cycle theory of surface metabolism. Microbiol Rev
52:452–484
▶ Chemolithoautotroph Wächtershäuser G (1990) Evolution of the first metabolic
▶ Isotopic Fractionation (Interstellar Medium) cycles. Proc Natl Acad Sci U S A 87:200–204
▶ Origin of Life
▶ Photoautotroph
▶ Rubisco
Autumnal Point
▶ Equinox
Alberts B, Bray D, Lewis J, Raff M, Roberts K, Watson JD
(1994) Molecular biology of the cell, 3rd edn. Garland,
New York
Berg IA, Kockelkorn D, Ramos-Vera WH, Say RF,
Zarzycki J, H€ugler M, Alber BE, Fuchs G (2010)
Available Water
Autotrophic carbon fixation in archaea. Nat Rev
Microbiol 8:447–460 ▶ Water Activity
Campbell NA, Reece JB (2002) Biology, 6th edn. Pearson,
Upper Saddle River
Ehrlich HL (2002) Geomicrobiology, 4th edn. Marcel
Dekker, New York
Evans MCW, Buchanan BB, Arnon DI (1966) A new Axial Tilt
ferredoxin-dependent carbon reduction cycle in a pho-
tosynthetic bacterium. Proc Natl Acad Sci U S
A 55:928–934
▶ Obliquity and Obliquity Variations
González-Toril E, Llobet-Brossa E, Casamayor EO,
Amann R, Amils R (2003) Microbial ecology of an
extreme acidic environment, the Tinto River. Appl
Environ Microbiol 69(8):4853–4865
Herter S, Fuchs G, Bacher A, Eisenreich WA
Azane
(2002) A bicyclic autotrophic CO2 fixation pathway
in Chloroflexus aurantiacus. J Biol Chem ▶ Ammonia
277:20277–20283
Ljundahl L, Irion E, Wood HG (1965) Role of corrinoids
in the total synthesis of acetate from CO2 by Clostrid-
ium thermoaceticum. Biochemistry 4:2771–2780
Madigan MT, Martinko JM, Parker J (2003) Brock biol- Azulmin
ogy of microorganisms, 10th edn. Pearson, Upper Sad-
dle River
Maier RM, Pepper IL, Gerba CP (2000) Environmental
▶ HCN Polymer
microbiology, 2nd edn. Academic, San Diego
B
Background Definition
Daniel Rouan Bacteria are a large group of single-celled phylo-

LESIA, Observatoire Paris-Site de Meudon, genetically related prokaryotes distinct from
Meudon, France ▶ Archaea. Bacteria have a wide range of shapes,
ranging from spheres to rods and spirals. They are
ubiquitous, growing in soil, water, extreme envi-
Definition ronments, and deep in the Earth’s crust (Vreeland
et al. 2000; Wanger et al. 2008). There are typi-
A background is a diffuse radiation field cally 40 million bacterial cells in a gram of soil
originating from no specific location in the sky. and a million bacterial cells in a milliliter of fresh
When observing a given field, it is that part of the water. There are approximately 5 1030 bacte-
signal which is not due to the objects of interest and ria on Earth, forming much of the world’s bio-
comes from extended or unresolved sources behind mass (Whitman et al. 1998). Bacteria are vital in
them. At millimeter wavelengths, the microwave recycling nutrients and important elements of
background, the fossil emission from the Big Bang, the geobiological cycles. Most have not been
is present in any direction of the sky. By extension, characterized, and only about half of the phyla
the molecular glow or the thermal infrared emis- of bacteria have species that can be grown in the
sion from the atmosphere as well as from the laboratory.
telescope that is superimposed on the astronomical Once regarded as plants constituting the
signal is also called background. class Schizomycetes, bacteria are now classified
as prokaryotes. Unlike the cells of animals
and other eukaryotes, bacterial cells do not
Bacteria contain a nucleus and rarely harbor membrane-
bound organelles. Although the term bacteria
Francisco Rodriguez-Valera traditionally included all prokaryotes, the
Microbiologia, Universidad Miguel Hernandez, scientific classification changed after the dis-
Campus San Juan, San Juan, Alicante, Spain covery in the 1970s that prokaryotes consist of
two very different groups of organisms
(domains) that evolved independently from
Keywords an ancient ▶ common ancestor. These evolu-
tionary domains are called bacteria and
Cell; Domain; Phylogeny; Prokaryote; Tree of life ▶ Archaea.
DOI 10.1007/978-3-662-44185-5
226 Bacteria
History known metabolic pathways and driving the func-

tioning of most ecosystems. The only known
Bacteria were first observed by Antonie van Leeu- metabolic strategy that is not found in bacteria
wenhoek in the seventeenth century, using a is the archaeal methanogenesis (DeLong and
single-lens microscope of his own design (Porter Pace 2001). Bacteria can actually close all bio-
1976). He called them “animalcules” and geochemical cycles on their own, so that a
published his observations in a series of letters to uniquely bacterial biosphere can be envisioned.
the Royal Society (van Leeuwenhoek 1684). The This is not true of either eukaryotes or Archaea
name bacterium was introduced much later, by (at least with the admittedly limited knowledge
Christian Gottfried Ehrenberg in 1838. that we have about this group).
Louis Pasteur demonstrated in the The classification of bacteria has undergone
mid-eighteenth century that ▶ fermentation is frequent changes in the last few years and is
caused by the growth of microorganisms and that subject to continuous controversy as is their
this growth is not due to spontaneous generation. relationship with the other major cellular types.
Along with his contemporary, Robert Koch, Pas- The widely accepted classification scheme is
teur was an early advocate of the germ theory of based on the 16S rRNA sequence comparisons.
disease. In his research into tuberculosis, Koch The most recent classification schemes
finally proved the germ theory, for which he was describe the following groups of Bacteria:
awarded a Nobel Prize in 1905. In Koch’s postu- ▶ Proteobacteria (Alpha-, Beta-, Gamma-,
lates, he set out criteria to test if an organism is the Delta-, Epsilonproteobacteria), Acidobacteria,
cause of a disease, and these postulates are still Aquificae, Chlorobi, Bacteroidetes, Chlamydiae/
used today. A major step in the study of bacteria Verrucomicrobia, Planctomycetes, Spirochaetes,
was the development, in the middle of the last Actinobacteria, Chloroflexi, Cyanobacteria,
century, of molecular biology techniques with Firmicutes, Tenericutes, Fusobacteria,
which much was learned about their biochemistry Synergistetes, Thermotogae, and Deinococcus/
and genetics. In 1977, Carl Woese (Woese and Thermus.
Fox 1977) made an important breakthrough in Some of these groups are extremely diverse in
our knowledge of evolution, when, in comparing metabolism, while others are very restricted.
the sequences of 16 S ribosomal RNA genes, he However, the availability of cultivated species,
observed that a group of prokaryotic organisms, properly studied for their physiology, is very
Archaea, have a separate line of evolutionary uneven (Rappé and Giovannoni 2003). At least
descent from that of bacteria (Woese et al. 1990). in some cases, this might influence our perception
of their true diversity. Cellular organization and
structure is also highly variable although most
Overview bacteria have a rigid cell wall consisting of one
single group of tridimensional polymers,
Bacteria is one of the two types of prokaryotic mureins, or peptidoglycans. Some bacteria,
organisms, the other is Archaea. Eukaryotic cells gram-negative bacteria, also have an extra mem-
actually represent a consortium of cells including brane outside the rigid polymer that provides a
a host or nucleus cytoplasm and the endosymbi- second permeability barrier. The outer mem-
otic organules: mitochondria and chloroplast, branes of the gram-negative bacteria have a
both of bacterial origin (Dyall et al. 2004). The very characteristic structure with a long chain
separation between bacteria and Archaea is polysaccharide facing the extracellular environ-
largely derived from molecular biology studies, ment and providing a hydrophilic envelope.
mostly on ▶ ribosome structure and components Some groups of gram-positive bacteria also
(proteins and ribosomal RNA) and ▶ transcrip- have an outer membrane, very different chemi-
tion machinery (RNA polymerase). Bacteria cally and more interlinked to the underlying poly-
indeed have representatives carrying out most mer. Sometimes these gram-positive bacteria
Bacteria 227
outer membranes are also quite as long as a high surface/volume ratio is

hydrophobic. Finally some bacteria have glyco- maintained.
protein S layers as rigid envelopes and some have Bacterial reproduction is asexual, by clonal
no rigid envelope at all. duplication of the cell, and originates large
▶ Motility is common in bacteria and is populations of clonal descent. This has been B
achieved by a characteristic structure, the bacte- used for pure culture isolation and study since
rial flagellum (Kojima and Blair 2004). This the origins of microbiology. However, bacteria
nanorotor is unique and different structurally have sex, often referred to as horizontal gene
and phylogenetically from its archaeal and transfer or lateral gene transfer. The impact of
eukaryotic counterparts. The archaeal flagellum this genetic exchange that has no part in repro-
is more similar to a different bacterial structure duction is considered fundamental in the evolu-
that is actually involved in some types of bacte- tion of bacteria. The absence of meiotic processes
rial motility by gliding. In spirochetes, the flexing and zygote formation leads to the possibility of
spiral body rotation is achieved by inward genetic exchange among very different partners.
directed flagella that are coiled over the cell. In this way, the barriers to genetic recombination
Some bacteria have other types of motility such in bacteria are very leaky, if present at all. The
as gliding over surfaces. This capacity is particu- impact of horizontal gene transfer in the evolu-
larly efficient in social bacteria that move in tion of the prokaryotic (bacterial and archaeal)
multicellular consortia. This is the case for world is still a matter of controversy, but some
myxobacteria (a group within the authors consider that a biologically consistent
deltaproteobacteria). tree of life based on evolutionary relationships
Although no organelles have been described could simply be a human fabrication.
in bacteria, there are examples of fairly complex
cellular structures, including the presence of
nuclear membranes, previously considered See Also
unique to eukaryotes (Fuerst 2005). Other cellu-
lar organule-like structures (Yeates et al. 2008) ▶ Aerobic Respiration
are magnetosomes, peroxisomes, gas vesicles, ▶ Anaerobic Respiration
and storage granules. Cell division in bacteria is ▶ Anoxygenic Photosynthesis
achieved by a cross-divisional cell wall septum, a ▶ Bioenergetics
constriction encompassing all the cell envelope ▶ Biofilm
layers. Although there is no mechanical cell ▶ Carbon Cycle, Biological
machinery such as the mitotic apparatus of ▶ Chemolithotroph
eukaryotes, bacteria have proteins homologous ▶ Chemoorganotroph
to actin that can produce mechanical modifica- ▶ Common Ancestor
tion of cell morphology (Shih and Rothfield ▶ Early Archean
2006). ▶ Fermentation
Cell shape and size varies widely in bacteria. ▶ Gram-Negative Bacteria
However, most bacterial cells conform to rela- ▶ Gram-Positive Bacteria
tively simple geometric shapes based in the cyl- ▶ Genotype
inder (rod), sphere (coccus), or the spiral coil ▶ Lateral Gene Transfer
(spirillum) (Young 2006). Cells can vary in size ▶ Metabolic Diversity
from fractions of a micron to many hundreds of ▶ Metabolism, Secondary
microns in giant bacteria such as Thiomargarita ▶ Microorganism
or Epulopiscium (Schulz and Jorgensen 2001). ▶ Motility
Bacterial size is limited by their osmotrophic ▶ Organelle
feeding, that is, their need to transport all nutri- ▶ Peroxisome
ents across the membrane. This is only efficient ▶ Phenotype
228 Bacterial Microcompartments
▶ Photosynthesis Whitman WB, Coleman DC, Wiebe WJ (1998) Prokary-

▶ Phylogenetic Tree otes: the unseen majority. Proc Natl Acad Sci USA
95:6578–6583. doi:10.1073/pnas.95.12.6578
▶ Phylogeny Woese C, Fox G (1977) Phylogenetic structure of the
▶ Phylum prokaryotic domain: the primary kingdoms. Proc Natl
▶ Prokaryote Acad Sci U S A 74(11):5088–5090. doi:10.1073/
▶ Proteobacteria pnas.74.11.5088
Woese CR, Kandler O, Wheelis ML (1990) Towards a
▶ Quorum Sensing natural system of organisms: proposal for the domains
▶ Respiration archaea, bacteria, and eucarya. Proc Natl Acad Sci U S A
▶ Ribosome 87:4576–4579. doi:10.1073/pnas.87.12.4576
▶ Sequence Analysis Xu J (2006) Microbial ecology in the age of genomics and
metagenomics: concepts, tools, and recent advances.
▶ Transcription Mol Ecol 15:1713–1731. doi:10.1111/j.1365-
▶ Translation 294X.2006.02882
Yeates TO, Kerfeld CA, Heinhorst S, Cannon GC, Shively
JM (2008) Protein-based organelles in bacteria:
References and Further Reading carboxysomes and related microcompartments. Nat
Rev Microbiol 6:681–691. doi:10.1038/nrmicro1913
DeLong E, Pace N (2001) Environmental diversity of Young K (2006) The selective value of bacterial shape.
bacteria and archaea. Syst Biol 50:470–478. Microbiol Mol Biol Rev 70(3):660–703. doi:10.1128/
doi:10.1080/106351501750435040 MMBR.00001-06
Dyall S, Brown M, Johnson P (2004) Ancient invasions: Zoetendal E, Vaughan E, de Vos W (2006) A microbial
from endosymbionts to organelles. Science world within us. Mol Microbiol 59:1639–1650.
304(5668):253–257. doi:10.1126/science.1094884 doi:10.1111/j.1365-2958.2006.05056
Fuerst J (2005) Intracellular compartmentation in
planctomycetes. Annu Rev Microbiol 59:299–328.
doi:10.1146/annurev.micro.59.030804.121258
Kojima S, Blair D (2004) The bacterial flagellar motor:
structure and function of a complex molecular
machine. Int Rev Cytol 233:93–134. doi:10.1016/
Bacterial Microcompartments
S0074-7696(04)33003-2
Porter JR (1976) Antony van Leeuwenhoek: tercentenary ▶ Carboxysomes, Structure and Function
of his discovery of bacteria. Bacteriol Rev 40:260–269
Rappé MS, Giovannoni SJ (2003) The uncultured micro-
bial majority. Annu Rev Microbiol 57:369–394.
doi:10.1146/annurev.micro.57.030502.090759
Schulz H, Jorgensen B (2001) Big bacteria. Annu Rev Bacterial Spore
Microbiol 55:105–137. doi:10.1146/annurev.
micro.55.1.105
▶ Endospore
Shih YL, Rothfield L (2006) The bacterial cytoskeleton.
Microbiol Mol Biol Rev 70(3):729–754. doi:10.1128/
MMBR.00017-06
van Leeuwenhoek A (1684) An abstract of a letter from
Mr. Anthony Leevvenhoek at Delft, dated 17 Sep
1683, containing some microscopical observations,
Bacteriochlorophyll
about animals in the scurf of the teeth, the substance
call’d worms in the nose, the Cuticula consisting of Juli Peretó
scales. Philos Trans (1683–1775) 14:568–574 Institut Cavanilles de Biodiversitat i Biologia
Vreeland R, Rosenzweig W, Powers D (2000) Isolation of
Evolutiva, Universitat de València, València,
a 250 million-year-old halotolerant bacterium from a
primary salt crystal. Nature 407:897–900. Spain
doi:10.1038/35038060
Wanger G, Onstott TC, Southam G (2008) Stars of the
terrestrial deep subsurface: a novel ‘star-shaped’ bac-
terial morphotype from a South African platinum
Synonyms
mine. Geobiology 6:325–330. doi:10.1111/j.1472-
4669.2008.00163 Chlorophylls
Banded Iron Formation 229
Definition
Baly’s Experiment
Bacteriochlorophylls are a family of
magnesium-porphyrin pigments present in Stéphane Tirard
anoxygenic photosynthetic bacteria and func- Centre François Viète d’Histoire des Sciences et B
tion both as light receptors and photochemical des Techniques EA 1161, Faculté des Sciences
reaction centers. et des Techniques de Nantes, Nantes, France
Definition
See Also
Regarding origin of life, Baly’s experiment (1922)
▶ Anoxygenic Photosynthesis
is one of the most emblematic organic chemistry
▶ Photosynthesis
experiments of the beginning of the twentieth cen-
tury (Baly 1871–1948). It has demonstrated that
ultraviolet light could act on a solution of water,
carbon dioxide, and ammonia and produce a lot of
Bacterirhodopsin organic compounds, such as sugars and amino acids.
J.B.S. Haldane quoted these experiments in his
Ricardo Amils famous text in 1929; for him, this was a chemical
Departamento de Biologia Molecular, proof of the possibility of a link between mineral
Universidad Autónoma de Madrid, Madrid, and organic chemistry in the prebiotic soup.
Spain
See Also
Definition
▶ Haldane’s Conception of Origins of Life
Bacteriorhodopsin is a membrane protein that
captures radiation energy which is used to create
a proton gradient. It can be found mainly in
Haloarchea. Bacteriorhodopsin is a membrane
Band Scan
protein usually found in two-dimensional crystal-
line patches known as purple membrane. The
▶ Molecular Line Survey
repeating element of the hexagonal lattice is com-
posed of three identical protein chains, each
rotated by 120 relative to the others. Each
chain has seven transmembrane alpha helices Banded Iron Formation
and contains one molecule of retinal buried
deep within. It is the retinal molecule that A. M. Mloszewska, Rasmus Nielsen Haugaard,
changes its conformation when absorbing a pho- Ernesto Pecoits and Kurt O. Konhauser
ton, resulting in a conformational change of the Department of Earth and Atmospheric Sciences,
bacteriorhodopsin promoting the proton pumping University of Alberta, Edmonton, AB, Canada
action. It can be considered the simplest photo-
synthetic system known. All other phototrophic
systems (bacteria, algae, chloroplasts) use chlo- Keywords
rophylls instead of bacteriorhodopsin to generate
proton gradients. Geobiology; Precambrian; Iron oxides; Chert
230 Banded Iron Formation
Synonyms Superior types. Algoma-type BIFs are compara-

tively small, with lateral extents rarely exceeding
BIF; Itabirite; Taconite 10 km, thicknesses ranging from 10 to 100 m, and
primary iron contents typically less than 1010 t
(e.g., Condie 1981). They are associated with
Definition volcanogenic complexes and are inferred to have
formed close to volcanic centers such as hydro-
A lithological term applied to a thinly bedded or thermal vents, back-arc basins, and intracon-
laminated chemical sedimentary rock consisting tinental rift zones. By comparison, Superior-type
of successive layers of fine-grained quartz, iron BIFs are hundreds of meters thick, have areal
oxides, carbonates, and/or silicates, typically extents to the order of 105 km2, and have total
containing 20–40 % iron and 40–50 % silica primary iron contents that exceed 1013 t. They
(James 1954; Trendall 2002; Klein 2005) (Fig. 1). are typically associated with sedimentary litholo-
gies (dolomite, quartzites, and shales) and are
inferred to have been deposited on the continental
Overview shelves and slopes of passive margins (Simonson
1985). Though the water depth of deposition is still
Banded iron formations (BIFs) comprise the larg- poorly constrained, the absence of wave- and
est iron resource on Earth. They formed through- current-generated features suggests a minimum
out much of the Precambrian (3,800–543 Ma), water depth of ~200 m (Trendall 2002).
reaching their maximum abundance between Banding in BIF is observed on a wide range of
2,700 and 2,400 Ma ago. Numerous examples scales, from coarse meter-thick macrobands and
can be found on almost every continent. Their centimeter-thick mesobands to millimeter- and
deposition has been linked to significant compo- submillimeter-scale bands. Among the latter is a
sitional changes in the Earth’s atmosphere and wide variety of varve-like repetitive laminae,
hydrosphere and possibly even to the diversifica- known as microbands, which may represent
tion of the biosphere. annual deposits (Trendall and Blockley 1970).
BIF has been classified on the basis of miner- Some Proterozoic iron formations are also
alogy, tectonic setting, and depositional environ- found to contain iron granules, oolites, and other
ment (Trendall 2002). The main iron mineral fragments embedded in a silica matrix (e.g., in the
phases were used to define four “iron formation Lake Superior Region and the Labrador Trough,
facies”: oxide, silicate, carbonate, and sulfide. The Canada; Nabberu Basin, Australia; Klein 2005).
dominant minerals, in their least metamorphosed This granular variety, called granular iron forma-
state, are hematite [Fe23+O3] and magnetite [Fe2 tion, possesses clear detrital textures and is
+
Fe23+O4] in the oxide facies, greenalite [(Fe2 thought to represent eroded and redeposited frag-
+
Mg)6Si4O10(OH)8] and minnesotaite ments of preexisting BIF (Trendall 2002).
[(Fe2+Mg)3Si4O10(OH)2] in the silicate facies, sid- A third type of BIF, the younger (750–560 Ma)
erite [Fe2+(CO3)] and ankerite [CaFe2+(CO3)2] in Rapitan type, is a special case in that it is linked to
the carbonate facies, and pyrite [Fe2+S2] in the global glaciations (“▶ snowball Earth” events).
sulfide facies (Klein 2005). Today, sulfide-facies The isolation of the oceans from the atmosphere
iron formations (i.e., pyritic carbonaceous shale or by worldwide glacial ice cover led to ocean stag-
slate) are no longer classified as BIF as they rep- nation and buildup of dissolved, hydrothermally
resent rock types that were deposited in different sourced Fe2+. As the ice melted and ocean circu-
environments and developed in sedimentary suc- lation reestablished itself, the iron became oxi-
cessions of different ages without systematic asso- dized and formed a suite of iron formations in
ciation with BIF. the oxic zone of upwelling areas (Klein 2005).
In terms of their size and lithological associa- Most Eo- and Mesoarchean BIFs belong to
tions, BIF are subdivided into Algoma and the Algoma-type and are associated with
Banded Iron Formation, Fig. 1 (a) Outcrop photo- core sample from the Joffre Member, Brockman Iron
graph of Algoma-type BIF with layers of magnetite BIF Formation, Hamersley Province, West Australia. Black
interbedded with felsic volcaniclastic rocks from the mesobands are composed mostly of dense magnetite, red
2.9 Ga Itilliarsuk BIF, West Greenland. (b) Outcrop pho- micro- and mesobands are composed of chert + hematite,
tograph of weakly folded magnetite-quartz BIF from the and gray micro- and mesobands are composed of
2.85 Ga Central Slave Cover Group, Slave Craton, North- chert + magnetite + Fe-rich carbonate (All photos cour-
west Territories, Canada. Brownish bands are secondary tesy of Rasmus Haugaard)
iron oxides and iron oxyhydroxides. (c) A pristine BIF
granite-greenstone belts (volcano-sedimentary Pilbara Craton (Hamersley Group, Western Aus-

sequences). They are found in the North Atlantic tralia), Kalahari Craton (Transvaal Supergroup,
Craton (Northern Labrador, southwestern Green- South Africa), São Francisco Craton
land), Guiana Shield (Venezuela, Guyana), (Quadrilátero Ferrı́fero), and the Superior Prov-
Kaapvaal Craton (South Africa and Swaziland), ince (Labrador Trough and Lake Superior
Liberian Shield (Sierra Leone, Guinea, Liberia, Regions, Canada/USA) (Trendall 2002; Klein
and Ivory Coast), and Yilgarn Craton (Western 2005). Major deposits of the Rapitan type include
Australia). Neoarchean to Proterozoic BIFs are the Rapitan Group in northern Canada and those
mostly of the Superior type and are found in the in the Urucum district of Brazil (Klein 2005).
Key Research Findings anomalies of Algoma-type BIFs compared to

their Superior-type counterparts suggest a higher
BIF Composition hydrothermal component in the former due to a
Iron formations can retain the chemical signa- higher input of high-temperature hydrothermal
tures of the seawater from which they precipi- fluids (>250 C) to Archean seawater caused by
tated provided that post-depositional element a hotter mantle (Arndt 1991). The redox-sensitive
mobility was minimal and may therefore be element Ce is used as a proxy for the redox state
used as potential proxies for ancient seawater of ancient seawater. Modern oxygenated seawa-
composition through time. Rare-earth element ter has a negative Ce anomaly due to oxidation of
and yttrium (REE + Y) profiles are extensively soluble Ce+3 to insoluble Ce+4 while the lack of
used to determine the origin of iron formations. Ce anomalies in Archean and early Paleopro-
Two aspects of the deviation of BIF REE + Y terozoic BIF REE + Y profiles suggests that
profiles from that of modern seawater, the Eu and they were instead precipitated from anoxic sea-
Ce anomalies of shale-normalized profiles, pro- water (e.g., Alexander et al. 2008 and references
vide key information about their depositional therein). Negligible fractionation of REE + Y
environments (Fig. 2). during the initial precipitation of the precursor
BIFs typically display positive Eu enrich- BIF iron oxyhydroxide minerals and their ability
ments which are thought to reflect the relative to withstand metamorphic conditions of up to
influence of hydrothermal fluids on the seawater amphibolite facies (Bau 1993) make these ele-
REE load at the time of deposition (Derry and ments an invaluable tool for deciphering the evo-
Jacobsen 1990). The more pronounced Eu lution of ancient seawater conditions.
Banded Iron Formation,

Fig. 2 Typical REE + Y
distribution in BIF showing
Eu and Ce anomalies as
depicted by average values
of BIF from the
ca. 3,700 Ga-old Isua
Supracrustal Belt (southern
West Greenland),
compared to the average
REE + Y signature of high-
temperature hydrothermal
fluids and average Pacific
seawater (Graph modified
after Alexander
et al. (2008); see references
therein)
To better understand the change in seawater signature as evidence of the direct carbonate pre-
chemistry through time and the effects of nutrient cipitation from an iron-rich, carbon-isotope-
limitations on early life, recent studies derived stratified water column with respect to dissolved
ancient seawater composition by applying inorganic carbon (e.g., Beukes and Klein 1990),
laboratory-derived partitioning coefficients petrographic evidence suggests that much of the B
between trace elements in seawater and iron carbonate mineral fraction in BIF is secondary,
oxyhydroxide particles to the absolute concentra- having grown during diagenesis (e.g., Ayres
tions of these elements in BIF (P, Bjerrum and 1972). More widely accepted opinions involve
Canfield 2002; Ni, Konhauser et al. 2009; Zn, microbially mediated precipitation of carbonate
Robbins et al. 2013). Because the partitioning of during burial diagenesis, including carbonate
P onto iron oxyhydroxides occurs in a predictable precipitation through microbially mediated
way, Bjerrum and Canfield (2002) used BIF to remineralization of (naturally isotopically light)
infer P concentrations in ancient seawater and its organic matter or due to the reduction of ferric
availability to ancient organisms. Konhauser iron and precipitation of carbonate minerals by
et al. (2009) showed that Ni concentrations means of the bacterial oxidation of methane (see
through time have changed drastically, with con- Bekker et al. 2010 and references therein).
centrations reaching up to 400 nM in the Archean The oxygen isotope (d18O) composition of
(compared to an average 9 nM in modern seawa- cherts in BIF is often used as a paleo-temperature
ter) due to an abundance of Ni-rich ultramafic proxy of the seawater from which they precipi-
rocks produced by a hotter mantle. Based on the tated because the oxygen isotope fractionation
Zn concentrations in BIF of various ages between silica and seawater is thought to be tem-
(an essential metal in the metabolism of eukary- perature dependent. Knauth and Lowe (2003)
otes), Robbins et al. (2013) suggested that Zn interpreted the depleted d18O composition of
concentrations in seawater have remained con- synsedimentary to very early diagenetic cherts
stant through time, contrary to previously in >3.2 Ga-old BIF (Barberton Greenstone Belt,
accepted ideas (e.g., Saito et al. 2003), forcing a South Africa) as indicative of high ocean temper-
reassessment of the influence of this metal on the ature values (55–85 C). This evidence was later
timing of eukaryote evolution. Future studies of corroborated by d18O data from an extensive
this kind on other bioessential trace metals will dataset of Archean to Phanerozoic cherts, propos-
advance our understanding of the timing and ing that sea surface temperatures decreased dras-
evolution of ancient microbial metabolisms. tically from about 70 C at 3,500 Ma ago to about
Yet another tool used to understand BIF gen- 20 C at 800 Ma ago (Robert and Chaussidon
esis and the environmental conditions under 2006). However, caution must be exercised
which they precipitated is stable isotopes. The when deriving paleo-seawater temperatures in
main traditional stable isotopes used in BIF stud- BIF whose origin is associated with
ies are carbon (d13C) and oxygen (d18O) while syndepositional hydrothermal alteration (e.g.,
nontraditional isotopes include iron (d56Fe), sili- BIF older than 3,500 Ma), since the derived tem-
con (d30Si), and most recently, chromium peratures will instead reflect local mixing condi-
(d53Cr), uranium (d238U), and germanium tions of hydrothermal fluids with overlying
(d74Ge). Carbon isotopes are primarily used as a seawater.
means of understanding the genesis of iron for- The study of iron isotopes (d56Fe) in BIF has
mations. Because the organic carbon fraction in gained momentum in recent years (e.g., Dauphas
BIF is so small (typically <0.5 wt.%; Gole and et al. 2004; Johnson et al. 2008). While the trace
Klein 1981), carbon-isotope studies tend to focus element distribution in BIF will be influenced by
instead on the carbonate mineral fraction. The post-depositional processes such as recrystalliza-
d13C signature in BIF is typically light, ranging tion and metasomatism, the iron isotope system
from +2.4 ‰ to 20.0 ‰ (Johnson et al. 2008). has been known to retain its pre-metamorphic
While one school of thought interprets this light signature despite such conditions (e.g., Dauphas
et al. 2004). Iron isotope values in BIF span the et al. 2009). It has been recently found that d238U
entire natural range (2.5 ‰ to +1.0 ‰), in con- in BIF and other marine sediments fractionate
trast to the near-constant d56Fe of igneous rocks differently under oxic and suboxic to euxinic
(~0 ‰) and other sedimentary rock types. They conditions, making it another novel geochemical
reflect a combination of three main processes: tracer of the Earth’s redox evolution (Weyer
(a) mineral-specific equilibrium fractionation, et al. 2008). Germanium in the oceans has a
(b) variation in the composition of fluids from short residence time (about 10,000 years). As
which they precipitated, and (c) the effects of Ge/Si ratios in seawater appear to vary in
bacterial Fe metabolic processing (Johnson response to climate, the d74Ge composition of
et al. 2003). Therefore, there are two important BIF and other marine sediments may offer new
applications of d56Fe in BIF: to provide informa- insight into the Earth’s climate history (Rouxel
tion on the abiotic and biogenic controls on et al. 2006).
ancient redox processes in the early oceans and
to determine the origins for heavily metamor- Controls on BIF Deposition
phosed rocks. Because the Fe contents of larger BIF is upward
The d30Si compositions of early diagenetic of 1013 t, the potential Fe source must be capable
cherts in BIF are often used to decipher the of supplying these vast quantities. A widely
genetic processes involved in their deposition accepted idea stems from classical BIF studies
and infer the ambient surface conditions on by Cloud (1973) and Holland (1973) which
early Earth (e.g., Andre et al. 2006). Typical established that the major components (Fe and
compositions which range from 2.5 ‰ to Si) in these rocks were derived from seawater.
0.5 ‰ reflect relative inputs by hydrothermal Additionally, certain aspects of the typical REE
(negative d30Si, e.g., Andre et al. 2006) and con- content in BIF led to the conclusion that these
tinental (positive d30Si, e.g. van den Boorn components were initially hydrothermally
et al. 2010) sources into the depositional basin. sourced and later mixed with overlying seawater
A predominantly hydrothermal silica source can (e.g., Graf 1978). The characteristic positive Eu
manifest itself as correlations in band-scale vari- anomalies found in BIF, a trait shared by modern
ations of Fe and Si isotopes, reflecting the dynam- hydrothermal vent fluids, is thought to indicate a
ics of the hydrothermal discharge (e.g., strong influence of hydrothermal fluids on the
ca. 2,700 Ma-old Wanderer BIF, Zimbabwe; BIF REE load and, by extension, Fe (Derry and
Steinhoefel et al. 2009). However, some of the Jacobsen 1990). This contrasting behavior of Eu
well-studied Superior-type BIFs whose silica to neighboring REE is thought to be related to the
precipitated from a well-mixed water column reduction of Eu3+ in hydrothermal solutions char-
far away from hydrothermal vents and continen- acterized by high temperatures (>250 C) and
tal drainage (e.g., on an isolated continental shelf low oxidation-reduction potentials. The generally
platform) show a uniform d30Si composition positive, mantle-like eNd values in BIF (where
within a single locality (e.g., the ca. 2,500 Ma- the eNd notation defines the departure of
143
old Transvaal BIF, South Africa, and Hamersley, Nd/144Nd from the Chondritic Uniform Reser-
Western Australia). voir evolution line), which contrast with the neg-
Analytical advances in stable isotope geo- ative eNd values of seawater, support the idea
chemistry have only recently allowed for high- that a large part of the Fe in BIF had been sourced
precision measurements of trace element isotopes from ancient submarine hydrothermal systems
such as Cr, U, and Ge in BIF. Chromium isotope (Jacobsen and Pimentel-Klose 1988).
(d53Cr) compositions in BIF can be used to inves- Our knowledge of the other major component
tigate the oxygenation of the ancient atmosphere, in BIF, silica, begins with the understanding that
where oxidative weathering will oxidize Cr(III) Precambrian seawater was likely close to super-
on land to the more mobile Cr(VI) which saturation with respect to amorphous silica until
becomes enriched in surface seawater (e.g., Frei the evolution of organisms which incorporate
Banded Iron Formation, Fig. 3 Three simplified the photic zone. (2) Fe(II) oxidation in an anoxic water
models for the oxidation of Fe2+ and the deposition of column by photoautotrophs such as green and purple
banded iron formations: (1) oxygenic photosynthesis and sulfur bacteria. (3) Abiotic oxidation of Fe2+ via
oxidation of Fe2+ by cyanobacterially produced oxygen in photooxidation
silica in their skeletons, such as diatoms and source since they show a perceptible hydrother-
radiolaria. The main sources of Si to the oceans mal signature in some BIFs (e.g., Steinhoefel
are hydrothermal fluids and continental runoff, et al. 2009), suggesting that the silica in BIF
which have compositionally distinct Ge/Si ratios may be derived from mixed hydrothermal and
that can be used to identify the origin of the silica continental sources.
in BIF. Hamade et al. (2003) established, using Prerequisites for BIF deposition necessitate
the Ge/Si ratios of chert bands in ca. 2,500 Ma not only a source of Si and soluble (reduced) Fe
BIF of the Dales Gorge Member (Hamersley but also a way to oxidize that Fe2+ into insoluble
Basin, Western Australia), that most of the silica Fe3+ (Fig. 3).
in BIF was continentally derived. Frei and Polat Traditional models of BIF deposition propose
(2007) later confirmed that this was also true for that Fe2+ was oxidized in the presence of free
Algoma-type BIFs, proposing that the spread in oxygen derived from oxygenic microbial photo-
Ge/Si ratios observed in the ca. 3,800 Ma-old Isua synthesis by cyanobacteria (Cloud 1965) and
Supracrustal Belt was due to the interaction of later through the direct utilization of O2 by aero-
hydrothermally iron-fertilized bottom waters bic chemolithoautotrophic bacteria (Holm 1989):
with silica-rich surface seawaters derived from
pre-4,000 Ma-old mafic land masses. Silicon iso-
topes have added some complexity to the issue of 2Feþ2 þ 1⁄2O2 þ 5H2 O ! 2FeðOHÞ3 þ 4Hþ
The signature feature of many iron formations

6Feþ2 þ 1⁄2O2 þ 16H2 O
is the distinctive banding made by alternating
! ½CH2 O þ 6FeðOHÞ3 þ 12Hþ
assemblages of silica/silicate and iron oxide min-
erals. Two fundamentally different models have
Indeed, the most voluminous BIFs (e.g.,
been proposed as to their formation. The first
Hamersley Group, Western Australia; Transvaal
suggests episodic pulsing of an iron-rich plume
Supergroup, South Africa) overlap in age with
into the shallow waters of a depositional basin.
the rise of atmospheric oxygen at ~2.4 Ga.
During periods of upwelling, increased bacterial
While this suggests that the Fe3+ component in
activity in the photic zone would induce the oxi-
these BIF formed via an oxic mechanism, the low
dation of Fe2+ to Fe3+, while periods of no
atmospheric oxygen concentrations in the
upwelling corresponded to low iron mineraliza-
Archean suggest that the pre-2,400 Ma BIF
tion and high silicification from background
formed via an anoxic mechanism. Both abiotic
waters (Morris 1993). Posth et al. (2008) demon-
and biogenic BIF formation mechanisms have
strated experimentally that banding could also be
been suggested. In the latter case, the oxidation
due to natural fluctuations in seawater tempera-
of Fe2+ into Fe3+ in an anoxic world can occur via
ture. In the summer, the warm waters promote
anoxygenic phototrophy, whereby some photo-
increased bacterial Fe2+ oxidation while
synthetic bacteria (e.g., green and purple sulfur
maintaining Si in solution, while during the win-
bacteria) can use Fe2+ as an electron donor for
ter, the decline in water temperature diminishes
carbon assimilation instead of water, thus pro-
bacterial activity but induces silicification. The
ducing Fe3+ instead of O2 (Garrels et al. 1973):
second view proposes that banding is the result of
post-depositional diagenetic processes in which
4Fe2þ þ CO2 þ 11H2 O Si was remobilized and segregated into bands
! CH2 O þ 4FeðOHÞ3 þ 8Hþ (Trendall and Blockley 1970). Significantly, cou-
pling the reduction of Fe3+ minerals to the oxida-
tion of organic matter not only explains the
The biogenic precipitation of iron oxides by sev-
reduced Fe mineralogy in BIF and the low
eral species of modern phototrophic bacteria has
organic-matter content but also explains the
been observed in both freshwater and marine
abundance of light C isotope signatures associ-
environments, and laboratory experiments dem-
ated with the interlayered carbonate minerals
onstrate that this form of metabolism could gen-
(Konhauser et al. 2005).
erate sufficient quantities of Fe3+ to account for
all the oxidized iron in BIF (Kappler et al. 2005).
BIF and Iron Ore
It has also been postulated that the absorption
Apart from being an important proxy for Precam-
of ultraviolet radiation by either Fe2+ or Fe(OH)+
brian seawater composition, BIFs are the main
in the water column could have triggered the
source of metallic iron for the steel industry.
hydrolyzation of these species according to the
Almost all iron extracted from BIF-hosted iron
following equation (Cairns-Smith 1978):
ore deposits are used to make pig iron (a mixture
of iron ore and coke produced in a blast furnace),
2Fe2þ ðaqÞ þ 2Hþ þ hv ! 2Fe3þ ðaqÞ þ H2 " which is the main component in the manufacture
of steel.
Although laboratory experiments have demon- BIF-hosted iron ore deposits can be divided
strated that this oxidative process could have into three classes on the basis of their iron con-
generated enough Fe3+ to account for all the ferric tent: (1) iron-rich primary BIFs containing
oxides in BIF, the process remains contentious as 30–45 wt.% Fe, (2) high-grade martite-goethite
the experiments were not done in solutions that ores with 56–63 wt.% Fe, and (3) high-grade
actually mimicked Precambrian seawater compo- hematite ores containing 60–68 wt.% Fe (Clout
sition (Konhauser et al. 2007). and Simonson 2005). The bulk of iron ore mined
Banded Iron Formation, Fig. 4 Sketch showing typical supergene enrichment of a magnetite-rich BIF leading to a
high-grade hematite-goethite ore deposit within the weathering zone (Modified from Clout and Simonson (2005))
today corresponds to high-grade iron deposits also presented. As observed in Table 1, the world
(class 1 and 2) formed by supergene iron enrich- output of iron is projected to increase from 2010
ment of precursor BIFs. The supergene enrich- to 2013 by about 13 %, that is, from 1.32 billion
ment process (Fig. 4) involves deep weathering metric tons (Gt) in 2010 to 1.52 Gt in 2013
from the downward movement of oxidizing (Menzie et al. 2013). The world iron resources
meteoric fluids that leach out the chert and car- (parts of the mineralization that is in such form
bonate components, leaving a residual accumu- and amount that economic extraction is not fea-
lation of oxidized and hydrous Fe3+ phases such sible) reached ca. 800 Gt of crude ore (defined as
as hematite, martite, and goethite (Webb it leaves the mine in an unconcentrated form)
et al. 2003; Clout and Simonson 2005). Martite containing 230 Gt of iron (US Geological Survey,
is often used as a textural term for hematite pseu- Mineral Commodity Summaries, January 2013).
domorphs after magnetite. The formation of these Superior-type BIF are the largest source of
high-grade iron ores took place long after depo- iron used in industry, where three of the largest
sition, likely during the Phanerozoic, when the deposits in the world are located in the Carajás
concentration of atmospheric oxygen, and thus and Quadrilátero Ferrı́fero iron provinces in Bra-
meteoric waters, was sufficient to oxidize precur- zil and the Hamersley iron province in Western
sor mineral phases such as magnetite (Fig. 4). Australia. These provinces account for close to
In some deposits, however, hydrothermal 40 % of the world’s iron reserves (Table 1). An
fluids facilitated the formation of high-grade example of the magnitude of these BIF deposits is
iron ore prior to supergene enrichment. Within seen within the Hamersley iron province where
the depositional basin, chert bands were hydro- the Paleoproterozoic Brockman Iron Formation
thermally replaced by carbonates, which in turn contains some of the largest single lithostra-
were dissolved and leached during later super- tigraphic BIF units known. These units are up to
gene enrichment (e.g., Tayor and Dalstra 2001). 360 m thick (e.g., the Joffre Member), having an
The world’s leading iron ore producers and areal extent of 105 km2 and estimated initial Fe
their historical and future estimated output of Fe content of 4.3 1013 t at the time of deposition
are shown in Table 1. (e.g., Trendall and Blockley 1970).
The iron reserves (part of the mineralization Table 1 shows that China is the largest pro-
that is economically feasible to extract at the time ducer and importer of iron in the world. The main
of determination) for each country as in 2011 are iron ore reserves of China, however, possess
Banded Iron Formation, Table 1 The world’s largest producers of iron 2000–2017 (estimated). See text for details.
Values extracted from Menzie et al. (2013) and US Geological Survey Mineral Commodity Summaries, January 2013
Reserves in billion
Fe content in thousand metric tons tons (Gt)
Average
ore Crude
grade (% ore Fe
Country Fe) 2000 2005 2010 2013 2015 2017 (2011) content
Australia 62 107,000 163,000 271,000 330,000 350,000 370,000 35,000 17,000
Brazil 66 141,000 186,891 247,772 250,000 250,000 260,000 29,000 16,000
Canada 64 22,700 19,333 23,300 28,000 28,000 30,000 6,300 2,300
China 64 73,600 134,000 350,000 410,000 420,000 430,000 23,000 7,200
India 64 48,600 97,500 166,000 170,000 172,000 174,000 7,000 4,500
Russia 58 50,000 56,100 58,500 59,000 59,500 60,000 25,000 14,000
South 62–65 21,570 24,900 36,900 46,700 48,900 49,500 1,000 650
Africa
Ukraine 55 30,600 37,700 43,000 45,000 48,000 50,000 6,000 2,100
USA – 39,703 34,202 32,000 32,000 32,000 32,000 6,900 2,100
World 607,000 837,000 1,320,000 1,520,000 1,660,000 1,750,000 170,000 80,000
relatively low-grade Fe content (~30 % Fe); conditions under which BIF formed. New
therefore, high export rates of high-grade iron BIF-based studies examining the paleoredox con-
ore are expected from China’s largest suppliers ditions a few million years prior to and after the
in the future (e.g., Australia and Brazil). great oxidation event (GOE, ca. 2.45 Ga) indicate
that this event likely occurred more gradually
than previously thought. As the products of the
Future Directions interplay between the mantle, the ocean, and the
biosphere, BIFs are important chemical archives
As marine chemical precipitates, BIFs hold great of ancient seawater composition. They not only
importance as proxies for ancient seawater chem- grant us an understanding into ancient mantle and
istry and, in this regard, may provide new insights surface process systematics but provide impor-
into the composition of the ancient marine bio- tant insights into the evolutionary pathways of
sphere and, ultimately, the atmosphere, through early life as well.
the biogenic gases that ancient plankton emitted.
Recent studies have begun to examine changing
Precambrian seawater chemistry in terms of See Also
bioessential trace metals, and the results obtained
from the BIF record are being corroborated by the ▶ Archean Environmental Conditions
record of these trace metals in marine black ▶ Archean Traces of Life
shales (e.g., Zn; Scott et al. 2012). Future work ▶ Bacteria
on BIF will undoubtedly investigate the temporal ▶ Earth’s Atmosphere, Origin and Evolution of
variations in more metals. Such geochemical ▶ Iron Cycle
studies will be enhanced by new analytical devel- ▶ Iron Isotopes
opments in trace metal stable isotopes, whereby ▶ Iron Oxides, Hydroxides and Oxy-hydroxides
relatively novel metal isotopes (e.g., d53Cr and ▶ Jaspilite
d238U) or multi-isotope surveys (e.g., combining ▶ Magnetite
d56Fe and d30Si; Steinhofel et al. 2009) will ▶ Ocean, Chemical Evolution of
improve our understanding of the paleoredox ▶ Oxygenation of the Earth’s Atmosphere
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KO, Beukes NJ, Gutzmer J (2010) Deciphering forma- ▶ Johnson UBV Bandpasses
tion processes of banded iron formations from the
Transvaal and the Hamersley successions by com-
bined Si and Fe isotope analysis using UV femtosec-
ond laser ablation. Geochim Cosmochim Acta
74:2677–2696
Taylor D, Dalstra H (2001) Genesis of high-grade hema- Barberton Greenstone Belt
tite ore bodies of the Hamersley Province, Western
Australia. Econ Geol 96:837–873 Nicholas Arndt
Trendall AF (2002) The significance of iron-formation in
the Precambrian stratigraphic record. Int Assoc
Sedimentol Spec Publ 33:33–66
Trendall AF, Blockley JG (1970) The iron formations of
the Precambrian Hamersley Group, Western Australia: Keywords
with special reference to associated crocidolite. West
Aust Geol Surv Bull 119:336
van den Boorn SHJM, van Bergen JM, Vroon PZ, de Vries Chert; Greenstone belt; Komatiite; Sedimentary
ST, Nijman W (2010) Silicon isotope and trace rocks; South Africa; Traces of life
Barberton Greenstone Belt 241
Definition 1980; de Wit et al. 1987; Dann 2000). The third

important development stemmed from the work on
The Barberton greenstone belt in South Africa is clastic and chemical sedimentary rocks. Following
one of the best-preserved successions of the recognition of the ancient age of the sedimen-
mid-Archean (3.57–3.21 Ga) supracrustal rocks tary sequences in the 1970s, several research B
in the world, together with the ▶ Pilbara Craton groups undertook a systematic search for micro-
in Western Australia. As such, it is a remarkable fossils in carbonaceous black ▶ cherts and shales.
natural laboratory where conditions and pro- Detailed sedimentological and geochemical stud-
cesses at the surface of the Archean Earth can ies followed and led to major advances in the
be studied in detail. The volcanic sequences understanding of surface processes on the early
include thick flows of ▶ komatiite, a type of Earth, including the observation of shallow-water
ultramafic lava named after the Komati River sedimentary rocks and silicified evaporites (Lowe
that flows through the belt, and records of large and Knauth 1977), sedimentary barite horizons
phreatomagmatic eruptions. Sedimentary (Heinrichs and Reimer 1977), seafloor alteration
sequences include ▶ cherts, ▶ banded-iron for- (Duchac and Hanor 1987), seafloor hot springs
mations, and barite whose compositions con- (de Ronde and Ebbesen 1996), semi-quantitative
strain the composition and temperature of evidence for tides (Eriksson and Simpson 2000), a
Archean oceans and preserve some of the earliest record of temperature of the Archean ocean
traces of life on Earth. The granitic rocks and (Knauth and Lowe 2003), and photosynthetic
metamorphic sequences provide information microbial mats (Tice and Lowe 2006). In addition,
about Archean tectonic processes. it was found that at least four sedimentary beds in
the Barberton belt contained sand-sized spherical
particles (spherules) interpreted to have formed by
History condensation of clouds of impact-generated rock
vapor and thus represent the oldest terrestrial
Geological mapping and research has been carried impact deposits (Lowe et al. 2003; Hofmann
out in the Barberton belt over much of the past et al. 2006).
century, but only in the 1970s did its geological
significance become apparent. Three develop-
ments were particularly important: The first was Overview
the development of increasingly accurate age dat-
ing that revealed the emplacement of volcanic and The Barberton greenstone belt is a small, cusp-
sedimentary rocks between 3.5 and 3.2 Ga ago shaped succession of volcanic and sedimentary
(e.g., Lopez Martinez et al. 1984; Armstrong rocks invaded on all sides by granitoid plutons or
et al. 1990). This discovery showed that the Bar- truncated by ductile shear zones. It is located
berton terrain, together with those of the Pilbara about 350 km east of Johannesburg and is world
Craton in Western Australia, contained the oldest famous for its komatiites, a type of ultramafic
known, well-preserved (i.e., metamorphosed to lava named after the Komati River that runs
only a low degree) volcanic and sedimentary through the southern part of the belt, and for
sequences. The second important discovery was thick sequences of sedimentary rocks, which
the recognition by Richard and Morris Viljoen of have yielded some of the earliest records of
the University of the Witwatersrand that many early life and of Earth’s early surface conditions.
of the ultramafic rocks of the sequence were The greenstone sequences, assigned to the ▶ Bar-
volcanic. They named this new rock type komatiite berton Supergroup, have been subdivided into
after the Komati River that flows through the belt. three stratigraphic units. From base to top, these
The geological and tectonic significance of these are (1) the Onverwacht Group, dominated by
rocks has been developed in numerous publica- ultramafic and mafic volcanic rocks; (2) the Fig
tions (e.g., Viljoen and Viljoen 1969; Anhaeusser Tree Group, a volcano-sedimentary succession
242 Barberton Greenstone Belt
made up of graywackes (a variety of sandstone depositional systems; the interface between vari-
with a clay-rich matrix), shales, cherts, and felsic ous regimes of erosion, transport, and deposition;
volcaniclastic rocks; and (3) the Moodies Group, the diagenetic processes near the surface; struc-
characterized by coarse-grained clastic sedimen- tural control on sedimentary composition, thick-
tary rocks, mainly sandstones and conglomerates. ness, and geometry; the thermal history and
Geological mapping has provided an increasingly evolution of sedimentary basins; and thus the
clearer picture of the detailed stratigraphy despite overall geodynamic setting on the early Earth.
the complex structure (Lowe et al. 2012). The The ultramafic lavas of the Barberton belt
protracted, 350-million-year-long evolution of the have unusual compositions that define the
region encompassed multiple tectonic events that Al-depleted or Barberton-type komatiite
include three or more cycles of volcanism and (Nesbitt and Sun 1976; Arndt et al. 2008).
sedimentation, deformation, and granite intrusion These rocks are formed through melting under
(Lowe and Byerly 1999a). This rich history led de unusual conditions in the mantle. Controversy
Wit et al. (1992) to use the region as the basis of surrounds the exact setting: most geologists sup-
their model for the formation of continental crust. port a model in which the melts form in an unusu-
Extensive field-based studies in the Barberton ally hot mantle plume (e.g., Arndt et al. 2008),
belt, starting from the early 1960s, provided evi- but others advocate melting in cooler conditions
dence for the existence, as early as 3.5 Ga, of a rich in an Archean subduction zone (e.g., Grove and
microbial ecosystem. Spherical, coccoidal, Parman 2004). Resolution of the issue has impor-
rod-shaped, and filamentous microscopic struc- tant implications for our understanding of
tures made up of carbonaceous matter and Archean geodynamics. Despite more than
interpreted as microfossils have been recorded 30 years of research, very few complete chemical
from Onverwacht and Fig Tree Group cherts. analyses of Barberton komatiites are available,
These rocks were deposited in shallow and deep yet this information is crucial if the potential of
marine environments, possibly in part on normal these rocks as tracers of Archean geodynamic
Archean oceanic crust and possibly associated with processes is to be realized. Black and white
hydrothermal activity (Walsh 1992; Altermann smokers on the Archean ocean floor, the exits of
2001; Westall et al. 2001; Tice and Lowe 2006). hydrothermal fluids that circulated through basal-
Domal stromatolites are also present in the Bar- tic crust, represent one possible setting for the
berton belt (Byerly et al. 1986), but their origin, emergence and evolution of life (Russell
together with other early Archean occurrences, is et al. 2005). Examples of these may well exist
controversial (Lowe 1994). More recently, Furnes in the Barberton greenstone belt, but, as for most
et al. (2004) reported micrometer-scale tubular of the crucial geological and biological aspects of
structures, interpreted to represent bioerosion fea- the Archean mentioned above, also these are the
tures, in the rims of Onverwacht Group pillow subject of considerable debate (e.g., de Ronde
basalts. Stable isotopic data have revealed the pos- and Ebbesen 1996; Lowe and Byerly 2007).
sible emergence of diverse groups of prokaryotes
including carbon-fixing Bacteria and Archaea,
methanogens, sulfate reducers, and possibly Future Directions
photosynthesizers. Structures reminiscent of mod-
ern microbial mats in shallow-water sandstones Much of future work in the belt will focus around
are widespread in parts of the Moodies Group scientific drilling projects that are aimed at recov-
(Heubeck 2009; Gamper et al. 2012). ering continuous sections of well-preserved vol-
The sedimentary sequences themselves can be canic and sedimentary rocks. Work on the
mapped in detail and measured at high strati- sedimentary sequences will provide information
graphic resolution. They thus provide abundant about erosion and sedimentation on the early
information about conditions at the Earth’s sur- Earth, the composition and temperature of
face: the spatial and temporal sequence of Archean seawater, and possible sites where life
Barberton Greenstone Belt 243
may have emerged and evolved. The study of de Ronde CEJ, Ebbesen TW (1996) 3.2 b.y. of organic
tidal sequences will provide information about compound formation near seafloor hot springs. Geol-
ogy 24:791–794
the dynamics of the Earth-Moon system, and the de Wit MJ, Hart RA, Hart RJ (1987) The Jamestown
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greenstone belt, South Africa. Geology 28:831–834
habitat of early life. Work on deeper sections Furnes H, Banerjee NR, Muehlenbachs K, Staudigel H, de
through the lower parts of the succession will Wit MJ (2004) Early life recorded in Archean pillow
provide information about tectonic processes lavas. Science 304:578–581
that operated during deposition of volcanic, sed- Gamper A, Heubeck C, Demske D, Hoehse M (2012)
Composition and microfacies of Archean microbial
imentary, and granitic rocks and during accretion mats (Moodies Group, Ca. 3.22 Ga, South Africa).
of these materials to the continent. In: Noffke N, Chafetz H (eds) Microbial mats in
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See Also Grove TL, Parman S (2004) Thermal evolution of the
Earth as recorded by komatiites. Earth Planet Sci Lett
▶ Archean Traces of Life 219:173–187
▶ Barberton Supergroup Heinrichs TK, Reimer TO (1977) A sedimentary barite
deposit from the Archean fig tree group of the Barber-
▶ Chert ton mountain land (South Africa). Econ Geol
▶ Impact Melt Rock 72:1426–1441
▶ Komatiite Heubeck C (2009) An early ecosystem of Archean tidal
▶ Pilbara Craton microbial mats (Moodies Group, South Africa, ca. 3.2
Ga). Geology 37:931–934
▶ Stromatolites Hofmann A, Reimold UW, Koeberl C (2006) Archean
spherule layers in the Barberton Greenstone Belt, South
Africa: a discussion of problematics related to the impact
References and Further Reading interpretation. In: Reimold WU, Gibson R (eds) Pro-
cesses on the early Earth, vol 405, Geological Society
Altermann W (2001) The oldest fossils of Africa – a brief of America special paper., pp 33–56
reappraisal of reports from the Archean. J Afr Earth Knauth LP, Lowe DR (2003) High Archean climatic tem-
Sci 33:427–436 peratures inferred from oxygen isotope geochemistry
Anhaeusser CR (1980) A geological investigation of the of cherts in the 3.5 Ga Swaziland supergroup, South
Archean granite-greenstone terrane south of the Africa. Geol Soc Am Bull 115:566–580
boesmanskop syenite pluton Barberton mountain Lopez Martinez M, York D, Hall CM, Hanes JA
land. Trans Geol Soc South Afr 9:73–106 (1984) Oldest reliable 40Ar/39Ar ages for terrestrial
Armstrong RA, Compston W, De Wit MJ, Williams IS rocks: Barberton mountainland komatiites. Nature
(1990) The stratigraphy of the 3.5–3.2 Ga Barberton 307:352–354
greenstone belt revisited: a single zircon ion micro- Lowe DR, Byerly GR (eds) (1999a) Geologic evolution of
probe study. Earth Planet Sci Lett 101:90–106 the barberton greenstone belt, South Africa. Geologi-
Arndt NT, Barnes SJ, Lesher CM (2008) Komatiite. Cam- cal Society of America special paper 329, p 312
bridge University Press, Cambridge Lowe DR, Byerly GR (2007) Ironstone bodies of the
Dann JC (2000) The komati formation, Barberton green- Barberton greenstone belt, South Africa: products of
stone belt, South Africa, part I: new map and magmatic a cenozoic hydrological system, not Archean hydro-
architecture. S Afr J Earth Sci 6:681–730 thermal vents! Geol Soc Am Bull 119:65–87
244 Barberton Greenstone Belt, Sedimentology
Lowe DR, Knauth LP (1977) Sedimentology of the Definition

onverwacht group (3.4 billion years), transvaal,
South Africa, and its bearing on the characteristics
and evolution of the early Earth. J Geol 85:699–723 ▶ Sedimentary rocks can be found throughout the
Lowe DR, Byerly GR, Kyte F, Shukolyukov A, Asaro F, ▶ Barberton Supergroup. In the volcano-
Krull A (2003) Spherule beds 3.47–3.24 billion years sedimentary Onverwacht Group, sedimentary
old in the Barberton greenstone belt, South Africa: a rocks make up less than 5 % of the succession.
record of large meteorite impacts and their influence
on early crustal and biological evolution. Astrobiology They consist of bedded ▶ cherts, representing a
3:7–48 variety of silicified clastic, chemical, and biogenic
Lowe DR, Byerly GR, Heubeck C (2012) Geologic map of sediments. Deep- to shallow-marine, fine-grained
the west-central Barberton Greenstone Belt, South siliciclastic sedimentary rocks make up the domi-
Africa, scale 1:25,000: Geol Soc America Map and
Chart Series No. 103 nant part of the Fig Tree Group. The Moodies
Nesbitt RW, Sun S-S (1976) Geochemistry of Archean Group is characterized by relatively coarse-
spinifex-textured peridotites and magnesian and grained, siliciclastic, alluvial, fluvial, shoreline,
low-magnesian tholeiites. Earth Planet Sci Lett and shallow-marine deposits. Detailed sedimento-
31:433–453
Russell MJ, Hall AJ, Boyce AJ, Fallick AE (2005) On logical and geochemical studies of the sedimen-
hydrothermal convection systems and the emergence tary rocks of the Barberton Supergroup have
of life. Econ Geol 100:419–438 provided major advances in the understanding of
Tice MM, Lowe DR (2006) The origin of carbonaceous surface processes in the Paleo- and Mesoarchean.
matter in pre-3.0 Ga greenstone terrains: a review and
new evidence from the 3.42 Ga buck reef chert. Earth
Sci Rev 76:259–300
Viljoen MJ, Viljoen RP (1969) Evidence for the existence Overview
of a mobile extrusive peridotitic magma from the
Komati Formation of the Onverwacht Group. Trans
Geol Soc S Afr Spec Publ 21:87–112 Bedded chert horizons of the Onverwacht Group
Walsh MM (1992) Microfossils and possible microfossils are typically 1–20 m thick and represent interflow
from the early Archean onverwacht Group, Barberton sedimentary units that were deposited on the sea-
mountain land, South Africa. Precambrian Res floor between phases of extrusive submarine vol-
54:271–293
Westall F, de Wit MJ, Dann JC, van der Gaast S, de Ronde canic activity. The cherts consist of a variety of
CEJ, Gerneke D (2001) Early Archean fossil bacteria silicified sediments (Lowe 1999; Tice and Lowe
and biofilms in hydrothermally influenced sediments 2006; Hofmann et al. 2013). Extensive early silic-
from the Barberton greenstone belt, South Africa. Pre- ification took place as a result of low-temperature
cambrian Res 106:93–116
hydrothermal activity on the seafloor, resulting in
excellent preservation of the sedimentary strata.
Silicified volcaniclastic sediments are common
and include silicified beds of ultramafic to mafic
Barberton Greenstone Belt, ash and accretionary lapilli. Laminated cherts of
Sedimentology various shades of gray to black represent mix-
tures of volcaniclastic material and carbonaceous
Axel Hofmann matter (Fig. 1). Deposition took place in
Department of Geology, University of low-energy, predominantly sub-wave base set-
Johannesburg, Auckland Park, Johannesburg, tings with episodic, high-energy current events
South Africa during which coarse-grained volcaniclastic mate-
rial and, in rare cases, meteorite impact ▶ ejecta
were deposited. Cherts have been used exten-
Keywords sively to study surface processes, seawater com-
position, and life in the Archean.
Archean Eon; Chert; Sedimentary rock; Exoge- The Fig Tree Group consists of a 2–3 km
nous; Onverwacht Group; Fig Tree Group; thick, largely siliciclastic, and volcaniclastic
Moodies Group sequence that is capped by felsic volcanic and
Barberton Greenstone Belt, Sedimentology 245
Barberton Greenstone
Belt, Sedimentology,
Fig. 1 Carbonaceous chert
with discontinuous layers
of silicified ultramafic ash
(gray chert) showing ripple B
lamination (characteristic
of subaqueous deposition).
Onverwacht Group,
Kromberg Formation, Farm
Josefsdal
volcaniclastic rocks (Heinrichs 1980; Lowe and few Ma under initially extensional and later com-
Nocita 1999; Hofmann 2005). In the southern pressional deformation (Heubeck et al. 2013).
part of the belt, a variety of siliciclastic
lithofacies with abundant felsic volcanic detritus
are present that formed in deep- to shallow-water,
fan delta, and alluvial environments (southern See Also
facies). The local presence of beds of jaspilitic
banded iron formation, chert, and barite indicates ▶ Archean Environmental Conditions
syndepositional hydrothermal activity. In the ▶ Barberton Greenstone Belt
northern part of the Barberton greenstone belt, ▶ Barberton Greenstone Belt, Traces of Early
the Fig Tree Group is mainly characterized by Life
turbiditic sandstones and shales that formed in ▶ Chert
relatively deepwater environment (northern ▶ Earth, Formation and Early Evolution
facies). Several spherule beds of quenched liquid ▶ Ejecta
silicate droplets in the Fig Tree Group represent ▶ Greenstone Belts
fallout and partially tsunami-reworked meteorite ▶ Sedimentary Rock
impact deposits (Lowe et al. 2003).
The ▶ Moodies Group consists of up to 3.7 km References and Further Reading
thick, quartz-rich, predominantly arenaceous
rocks, in contrast to the quartz-poor and matrix- Anhaeusser CR (1976) The geology of the Sheba hills area
rich Fig Tree graywackes. It consists of alluvial of the Barberton mountain land, South Africa, with
conglomerate, braided fluvial, tidal and shallow- particular reference to the Eureka syncline. Trans
Geol Soc S Afr 79:253–280
marine sandstones, and minor siltstone, jaspilite, Eriksson KA, Simpson EL (2000) Quantifying the oldest
and banded iron formation (Anhaeusser 1976; tidal record: the 3.2 Ga Moodies Group, Barberton
Heubeck and Lowe 1994). Intertidal sedimentary greenstone belt, South Africa. Geology 28:831–834
rocks are locally well preserved. Tidal bundles Heinrichs T (1980) Lithostratigraphische Untersuchungen
in der Fig Tree Gruppe des Barberton Greenstone Belt
and microbial mat-related sedimentary structures zwischen Umsoli und Lomati (S€ udafrika). Göttinger
have been reported (Heubeck and Lowe 1994; Arb Geol Paläont 22:118
Eriksson and Simpson 2000). Moodies strata Heubeck C, Lowe DR (1994) Depositional and tectonic
may include some of the oldest preserved aeolian setting of the Archean Moodies Group, Barberton green-
stone belt, South Africa. Precambrian Res 68:257–290
deposits (Simpson et al. 2012). Deposition of the Heubeck C, Engelhardt J, Byerly GR, Zeh A, Sell B,
Moodies Group was syntectonic with final green- Luber T, Lowe DR (2013) Timing of deposition and
stone belt deformation and occurred within only a deformation of the Moodies Group (Barberton
246 Barberton Greenstone Belt, Traces of Early Life
greenstone belt, South Africa): very-high-resolution of Synonyms

Archaean surface processes. Precambrian Res
231:236–262
Hofmann A (2005) The geochemistry of sedimentary Early Archean; Early life
rocks from the Fig Tree Group, Barberton greenstone
belt: implications for tectonic, hydrothermal and sur-
face processes during mid-Archaean times. Precam- Definition
Hofmann A, Bolhar R, Orberger B, Foucher F (2013)
Cherts of the Barberton greenstone belt: petrology The Barberton Greenstone Belt of South Africa
and trace-element geochemistry of 3.5 to 3.3 Ga old and Swaziland, together with the greenstone belts
silicified volcaniclastic sediments. South African Jour- of the Pilbara region in Australia, hosts the oldest
nal of Geology 116:297–322
Lowe DR (1999) Petrology and sedimentology of cherts (3.5–3.3 Ga) well-preserved rocks containing
and related silicified sedimentary rocks in the Swazi- fossil signatures of life. These traces occur in
land supergroup. In: Lowe DR, Byerly GR (eds) Geo- thin layers of volcaniclastic sediments,
logic evolution of the Barberton greenstone belt, sandwiched between thick successions of mafic
vol 329. Geol Soc Am Spec Pap, South Africa,
pp 83–114 to ultramafic volcanic lavas. The remains of
Lowe DR, Nocita BW (1999) Foreland basin sedimenta- microbial mats are relatively common in
tion in the Mapepe formation, southern-facies Fig tree shallow-water deposits, while deeper-water
group. In: Lowe DR, Byerly GR (eds) Geologic evo- deposits contain reworked, detrital remains of
lution of the Barberton greenstone belt, vol 329. Geol
Soc Am Spec Pap, South Africa, pp 233–258 organic matter and microbial mats. Somewhat
Lowe DR, Byerly GR, Kyte F, Shukolyukov A, Asaro F, younger clastic sediments of the 3.2 Ga-old
Krull A (2003) Spherule beds 3.47–3.24 billion years Moodies Group have yielded relatively large cel-
old in the Barberton Greenstone Belt, South Africa: a lular remains of possible planktonic organisms
record of large meteorite impacts and their influence
on early crustal and biological evolution. Astrobiology and abundant microbial mats.
3:7–48
Simpson EL, Eriksson KA, Muller WU (2012) 3.2 Ga
eolian deposits from the Moodies Group, Barberton Overview
greenstone belt, South Africa: implications for the
origin of first-cycle quartz sandstones. Precambrian
Res 214–215:185–191 The early to mid-Archaean (3.5–3.2 Ga) Barber-
Tice MM, Lowe DR (2006) The origin of carbonaceous ton Greenstone Belt consists of kilometer-thick,
matter in pre-3.0 Ga greenstone terrains: a review and largely subaqueous volcanic rocks and their ero-
new evidence from the 3.42 Ga Buck Reef Chert. Earth
Sci Rev 76:259–300 sional products and of sedimentary rocks includ-
ing Banded Iron Formations (BIFs), cherts, shales,
barite, siltstones, sandstones, and conglomerates.
Largely basic to ultrabasic lavas were extruded
Barberton Greenstone Belt, Traces of onto relatively shallow-water platforms. Inter-
Early Life spersed with the volcanics are thin horizons of
volcaniclastic and hydrothermal sediments depos-
Frances Westall ited in water depths ranging from littoral to
Centre de Biophysique Moléculaire, CNRS, sub-wave base. Early diagenetic silicification of
Orléans Cedex 2, France sediments and underlying volcanics was related to
circulating hydrothermal fluids (Hofmann and
Bohlar 2007) and seawater oversaturation with
Keywords respect to silica (Lowe and Byerly 1986). The
cherts and unsilicified siliciclastic sediments host
Early Archaean; Biosignatures; Early life; Pro- a variety of traces of early life.
karyotes; Microfossils; Carbon isotopes; Basalt; Early studies reported the occurrence of pos-
Volcanic sediments; Anaerobic photosynthesis; sible cellular microfossils (e.g., Knoll and
Chemolithotrophy Barghoorn 1977). Microbial corrosion features,
Barberton Greenstone Belt, Traces of Early Life 247
Barberton Greenstone Belt, Traces of Early Life, filamentous microbial mat from ~3.4 Ga-old cherts from
Fig. 1 (a) Purported microbial tunnels in the vitreous Barberton (Walsh 2004). (d) Well-preserved silicified fil-
rind of an ~3.5 Ga-old pillow lava from the Barberton aments in a 3.3 Ga-old microbial mat from Barberton
Greenstone Belt (Furnes et al. 2004). (b) Silicified colony (Westall et al. 2006b). (e) Acritarch (organic-walled ves-
of chemolithotrophic microorganisms on the surface of a icle of unknown biological affinity) from 3.2 Ga-old
volcanic particle in silicified volcaniclastic sediments siliciclastic sediments from the Moodies Group, Barber-
from the Pilbara, Australia (Westall et al. 2006a). (c) ton (Javaux et al. 2010)
Thin-section micrograph of possible photosynthetic
typical of those produced by chemolithotrophic confirmed (McLoughlin et al. 2012; Grosch

microorganisms in the vitreous rinds of pillow et al. 2014). The hollow structures in the surfaces
lavas, were described by Furnes et al. (2004, of the pillow basalts are lined by Ti-oxides that
2007) from the Kromberg Formation contain early sulfide inclusions showing d 34S
(ca. 3.4 Ga), but their biogenicity has yet to be depletions (8–45 ‰) which is consistent with an
248 Barberton Greenstone Belt, Traces of Early Life
early microbial origin for the sulfides. No micro- the presence of a variety of microbial organisms
fossils are associated with these tunnels. Fossil- including photosynthesizers (pelagic as well as
ized colonies of chemotrophic organisms that benthonic), heterotrophs such as methanogens,
could have produced such features have also and chemotrophs (see review in Westall 2011).
been described in similarly aged strata Younger littoral sediments from the 3.2 Ga-old
(3.446 Ga) (Westall et al. 2006a, 2011a) from Moodies Group have revealed a wealth of fossil
the Pilbara, Australia (Fig. 1). microbial structures. Crinkled, domed, and anas-
The surfaces of volcaniclastic sediments tomosing photosynthetic microbial mats occur on
deposited in shallow-water environments were the surfaces of coarse-grained, tidal-zone sand-
host to microbial mats and biofilms which prob- stones (Noffke et al. 2006; Heubeck 2009). Cor-
ably formed by anaerobic photosynthetic organ- relative siltstones have yielded compressed
isms. When observed in petrographical thin remains of large (up to 300 mm diameter) organic,
sections, they are characterized by packets of hollow, spherical structures (acritarchs)
fine-grained, carbon-rich layers that form more interpreted as unicellular microorganisms or colo-
or less continuous wispy, wavy horizons on sed- nial envelopes (Javaux et al. 2010), thus
iment surfaces (Walsh 1992, 2004; Walsh and documenting the existence of a diverse biota.
Lowe 1999; Tice and Lowe 2004, 2006; Tice
2009). Tice (2009) demonstrated that environ-
mental factors, such as current energy, controlled See Also
the style of microbial mats preserved in the
3.42 Ga-old Buck Reef Chert. Formed at water ▶ Anoxygenic Photosynthesis
depths between storm base and fair-weather wave ▶ Archean Traces of Life
base, anastomosing and mesh-like mats occur in ▶ Barberton Greenstone Belt
sediments consisting of coarse-grained fluffy car- ▶ Biomarkers
bonaceous grains, while finely laminated mats ▶ Chemolithotroph
occur in finer-grained sediments deposited ▶ Microbial Mats
under quiet conditions. ▶ Microfossils
Rarely, carbonaceous filaments are observed ▶ Pilbara Craton
in silicified interstitial spaces between mat layers
(Walsh 1992). Although the organisms forming
the mats are seldom preserved, a scanning elec- References and Further Reading
tron microscope study of an extremely well-
preserved silicified biofilm documented the Furnes H, Banerjee NR, Muehlenbachs K, Staudigel H, de
Wit M (2004) Early life recorded in Archean pillow
presence of filaments 0.25 mm in diameter and lavas. Science 304:578–581
tens of microns in length embedded in chert Furnes H, Banerjee NR, Staudigel H, Muehlenbachs K,
interpreted as thick polymer on the mat surface McLoughlin N, de Wit M, van Kranendonk M (2007)
(Westall et al. 2006b). Detailed in situ morpho- Comparing petrographic signatures of bioalteration in
recent to Mesoarchean pillow lavas: tracing subsurface
logical and geochemical analysis of this particu- life in oceanic igneous rocks. Precambrian Res
lar biofilm demonstrated that the biofilm had 158:156–176
been undergoing incipient calcification, probably Grosch EG, McLoughlin N, Lanari P, Erambert M, Vidal
due to the action of sulfate-reducing microorgan- O (2014) Microscale mapping of alteration conditions
and potential biosignatures in basaltic-ultramafic rocks
isms degrading the lower, dead layers of the on early Earth and beyond. Astrobiology 14:216–228
biofilm prior to early silicification that preserved Heubeck C (2009) An early ecosystem of Archean tidal
it in three dimensions (Westall et al. 2011b). microbial mats (Moodies Group, South Africa, ca. 3.2
The silicified sediments also contain detrital Ga). Geology 37:931–934
Hofmann A, Bolhar R (2007) The origin of carbonaceous
particles of carbon that may be derived from cherts in the Barberton Greenstone Belt and their sig-
microbial colonies. Thus, carbon and sulfur iso- nificance for the study of early life in mid-Archaean
tope signatures have been interpreted to indicate rocks. Astrobiology 7(2):355–388
Barberton Supergroup 249
Javaux EJ, Marshall CP, Bekker A (2010) Organic-walled Westall F, Foucher F, Cavalazzi B, de Vries ST,
microfossils in 3.2-billion-year-old shallow-marine Nijman W, Pearson V, Watson J, Verchovsky A,
siliciclastic deposits. Nature 463:934–938 Wright I, Rouzaud JN, Marchesini D, Anne S
Knoll AH, Baghoorn ES (1977) Archean microfossils (2011a) Early life on Earth and Mars: a case study
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South Africa. Science 198:396–398 Planet Space Sci 59:1093–1106 B
Lowe DR, Byerly GR (1986) Archean flow-top alteration Westall F, Cavalazzi B, Lemelle L, Marrocchi Y, Rouzaud
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McLoughlin N, Grosch EG, Kilburn MR, Wacey D (2012) Thiel V, Southam G, MacLean L, Wirick S,
Sulfur isotope evidence for a Paleoarchean subseafloor Hofmann A, Meibom A, Robert F, Défarge
biosphere, Barberton, South Africa. Geology C (2011b) Implications of in situ calcification for pho-
40:1031–1034 tosynthesis in a ~3.3 Ga-old microbial biofilm from the
Noffke N, Eriksson K, Hazen RM, Simpson EL (2006) Barberton greenstone belt, South Africa. Earth Planet
A new window into early Archean life: microbial mats Sci Lett 310:468–479
in Earth’s oldest siliciclastic tidal deposits (3.2 Ga
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Axel Hofmann
Tice M, Lowe DR (2004) Photosynthetic microbial mats
in the 3,416-Myr-old ocean. Nature 431:549–552 Department of Geology, University of
Tice MM, Lowe DR (2006) The origin of carbonaceous Johannesburg, Auckland Park, Johannesburg,
matter in pre-3.0 Ga greenstone terrains: a review and South Africa
new evidence from the 3.42 Ga Buck Reef Chert. Earth
Sci Rev 76:259–300
Walsh MM (1992) Microfossils and possible microfossils
from the early Archean Onverwacht Group, Barberton Synonyms
Mountain Land, South Africa. Precambrian Res
54:271–293
Swaziland Supergroup (outdated)
Walsh MM (2004) Evaluation of early Archean
volcaniclastic and volcanic flow rocks as possible
sites for carbonaceous fossil microbes. Astrobiology
4:429–437 Definition
Walsh MM, Lowe DR (1999) Modes of accumulation of
carbonaceous matter in the early Archaean: a petro-
graphic and geochemical study of carbonaceous cherts The volcano-sedimentary sequence that forms
from the Swaziland Supergroup. In: Lowe DR, Byerly the Barberton greenstone belt has been grouped
GR (eds) Geologic evolution of the Barberton green- stratigraphically as the Barberton Supergroup,
stone belt, South Africa. Geological Society of Amer-
renamed from the Swaziland Supergroup of
ica Special Paper, 329, Boulder, Colorado, pp 115–132
Westall F (2011) Early life. In: Gargaud M (ed) Origins of older literature. The Barberton Supergroup
life, an astrobiology perspective. Cambridge Univer- formed ca. 3.55–3.22 Ga ago and is subdivided
sity Press, Cambridge, pp 391–413 in ascending order into three major stratigraphic
Westall F, de Vries ST, Nijman W, Rouchon V, Orberger B,
units: (1) The Onverwacht Group dominantly
Pearson V, Watson J, Verchovsky A, Wright I, Rouzaud
J-N, Marchesini D, Anne S (2006a) The 3.466 Ga consists of submarine ultramafic-mafic volcanic
Kitty’s Gap Chert, an early Archaean microbial ecosys- rocks and minor felsic volcanic and silicified
tem. In: Reimold WU, Gibson R (eds) Processes on the ▶ sedimentary rocks. (2) The Fig Tree Group is
early Earth, vol 405, Geological Society of America
comprised of shale, graywacke, and felsic
Special Paper. Geological Society of America, Boulder,
pp 105–131 volcaniclastic rocks with minor conglomerate,
Westall F, de Ronde CEJ, Southam G, Grassineau N, chert, baryte, and banded iron formation.
Colas M, Cockell C, Lammer H (2006b) Implications (3) The Moodies Group consists of shallow
of a 3.472–3.333 Ga-old subaerial microbial mat from
marine to fluvial sandstone and conglomerate
the Barberton greenstone belt, South Africa for the UV
environmental conditions on the early Earth. Philos with minor felsic and mafic volcanics, shale,
Trans R Soc London Ser B 361:1857–1875 and banded iron formation. These groups have
250 Barophile
been subdivided in to a number of formations purposes of most calculations, the system can be
which differ in lithology, thickness, and facies considered to be concentrated at the position of
across the major fault zones that separate the the barycenter, with a total mass equal to the sum
Barberton greenstone belt into discrete fault- of the masses of the individual objects. The
bounded segments. The excellent degree of pres- barycentric radial velocity of an object, such as
ervation and exposure, continuous outcrop of a star, as measured by a terrestrial observer is
mostly subvertically dipping strata, and the high calculated relative to the center of mass of the
lithologic variability made possible the detailed Solar System. This is slightly different from the
mapping of the units of the Barberton Super- heliocentric velocity of such a star, which is cen-
group. This in turn provided the base for targeted tered on the Sun and varies slightly with time
sampling and major contributions to the under- according to the position of the planets (since
standing of the Archean. the motion of the planets produces slight changes
in the Solar System’s barycenter).
See Also
See Also
▶ Archean Environmental Conditions
▶ Barberton Greenstone Belt ▶ Astrometric Planets
▶ Barberton Greenstone Belt, Sedimentology
▶ Barberton Greenstone Belt, Traces of Early
Life
▶ Earth, Formation and Early Evolution Barycenter Velocity
▶ Greenstone Belts
▶ Center of Mass Velocity
Barophile
Barite
▶ Piezophile
Christoph Heubeck
Institut f€ur Geowissenschaften, Friedrich-
Schiller-Universität Jena, Jena, Germany
Barycenter
David W. Latham Definition

Harvard-Smithsonian Center for Astrophysics,
Cambridge, MA, USA Barite (also Baryte), BaSO4, is a widespread,
usually white or colorless mineral which can be
deposited by biogenic, hydrothermal, and evapo-
Synonyms ritic processes. In sedimentary environments, the
main sources of barium ions are submarine
Center of mass hydrothermal vents; the sulphate ion is common
in Phanerozoic seawater.
The origin of primary stratiform bladed baryte
Definition crystals on Archean sea floors (“crystal lawns”) is
debated because the highly oxidized sulfate ion is
In astronomy, the barycenter is the center of mass generally thought to have been in short supply in
of a system of two or more bodies. For the Archean oceans. However, some of these barite
Basic and Acid Rock 251
deposits may have formed from sulfate which had

been generated photolytically as aerosols in the Basaltic Flood Plains
Archean atmosphere and then washed into the
oceans. Others may be late-diagenetic pseudo- ▶ Mare, Maria
morphs after calcium sulphate. Studies on ▶ Trapps B
S isotopes indicate that some barite played a
role in bacterial sulphate reduction, one of the
earliest metabolic processes.
Base Pair
Juli Peretó
Basalt Institut Cavanilles de Biodiversitat i Biologia
Evolutiva, Universitat de València, València, Spain
Nicholas Arndt
Definition
Definition Base pair is a pair of nucleic acid bases each in a

different nucleotide monomer in the same
Basalt is a fine-grained, dark-colored mafic volca- (intramolecular) or different (intermolecular)
nic rock composed of plagioclase, ortho- or ▶ nucleic acid strands and linked to one another
clinopyroxene, and minor Fe-Ti oxides, with or by specific hydrogen bonds. The canonical base
without olivine. Porphyritic samples contain large pairs in ▶ DNA (Watson-Crick pairs) contain one
crystals (phenocrysts) of olivine, pyroxene, or pla- purine and one pyrimidine in antiparallel posi-
gioclase dispersed in a fine-grained glassy matrix, tions: adenine binds thymine in DNA – uracil in
also called groundmass. Gas-rich samples contain RNA (through two hydrogen bonds) – and gua-
abundant vesicles. Basalt contains 45–52 % SiO2 nine binds cytosine (through three hydrogen
and 40–90 % ferromagnesian minerals. Basalt bonds). There are also many examples of
erupts as pillow lava, thick sheet flows, or frag- non-Watson-Crick pairing, especially in three-
mental scoria. It is the most common rock of the dimensional structures of RNAs.
Earth’s ▶ oceanic crust and in lunar maria (ancient
flood-basalt plains corresponding to the dark sur-
faces of the ▶ Moon). Basalt is also present in the See Also
crust of ▶ Mars and ▶ Venus. It forms by partial
melting of the mantle and erupts in diverse tec- ▶ Anticodon
tonic settings: mid-ocean ridges, oceanic islands, ▶ Codon
subduction zones, continental rifts, and volcanic ▶ DNA
plateaus. ▶ Genetic Code
▶ Nucleic Acids
See Also ▶ RNA
▶ Igneous Rock
▶ Mafic and Felsic
▶ Mars
▶ Moon, The Basic and Acid Rock
▶ Oceanic Crust
▶ Venus ▶ Mafic and Felsic
252 Bathybius Haeckelii
of the ▶ European Space Agency’s Mars Express

Bathybius Haeckelii mission. It was launched in June 2003 and was
supposed to land in December 2003. Unfortu-
Stéphane Tirard nately, all contact with the 33.2 kg lander was
Centre François Viète d’Histoire des Sciences et lost a few days before touchdown, and its fate
des Techniques EA 1161, Faculté des Sciences et remains a mystery. The lander was a compact
des Techniques de Nantes, Nantes, France assemblage of instruments designed to address sci-
ence questions ranging from the search for traces of
past and extant life, measurement of the composi-
Definition
tion of the atmosphere and other environmental
parameters, the oxidation state at the surface, and
In June 1857, the Britannic ship “The Cyclops”
analysis of the geomorphology of the landing site
found a very special matter on the bottom of the
in a sedimentary Basin of Isidis Planitia.
North Atlantic Ocean. Some chemists and biolo-
gists considered that is was a very simple living
matter and thought that it resolved the problem of
Overview
the origin of life. Indeed, according to them, it
constituted an example of spontaneous genera-
The lander was constructed by a British consortium,
tion and a link between inert matter and living
coordinated by Prof. Colin Pillinger of the Open
matter. Thomas Huxley (1825–1895) himself
University. It was equipped with a 75 cm long
named it Bathybius haeckelii. However, in
robotic arm holding at its end an array of instru-
1876, a chemist revealed that it was calcium
ments called the PAW (Payload Adjustable Work-
sulfate and not living matter.
bench) that included stereo cameras, a Mössbauer
Discussions about Bathybius took place dur-
spectrometer to measure the oxidation states of
ing the period of the debate about spontaneous
iron-containing compounds, an X-ray spectrometer
generations. Therefore, during few years the pos-
for determining mineral composition, and a
sibility of the production of this matter, very
(dentist’s) drill for collecting samples. Also
closed to living matter, was highly considered
onboard the lander was a gas analysis package
by many chemists and biologists.
(GAP) that included a gas chromatograph-mass
spectrometer to analyze carbon isotopes as a signa-
See Also ture for life. Finally, a “mole” or subsurface sam-
pler (PLUTO – Planetary Undersurface Tool) was
▶ Huxley’s Conception on Origins of Life designed to penetrate beneath the loose regolith
▶ Protoplasmic Theory of Life surface to obtain samples for analysis.
The 1 m diameter lander was equipped with a
UHF radio antenna, telecommunications, a bat-
tery, electronic processors, heaters, solar panels,
Beagle 2 and other payload instruments, such as radiation
and oxidation sensors to address the environmen-
Frances Westall tal objectives of the mission. Upon arrival, the
Centre de Biophysique Moléculaire, CNRS, lander was supposed to have broadcast a piece of
Orléans, Cedex 2, France music specially composed by the British rock
band Blur.
Various hypotheses involving malfunctioning
Definition of various pieces of equipment have been put
forward to explain the failure of the mission.
The Beagle 2 lander, named after the famed ship in The lander could have bounced off the atmo-
which Charles Darwin traveled the world, was part sphere of Mars and burned up, or the parachute
Benthic Mats 253
Belcher Group,
Microfossils, Fig. 1 The
mat-building colony of the
cyanobacteria
Eoentophysallis
belcherensis, preserved in B
stromatolites of the 1.5 Ga
Bil’yakh Group, Siberia
(Photograph courtesy of
A. Knoll)
failed to deploy and/or the airbags did not func- 1.9 Ga. These rocks include ▶ chert lenses and
tion and the lander crashed onto the Martian nodules in silicified ▶ stromatolites growing
surface, or perhaps the backshell became in tidal and shallow subtidal waters on a carbon-
entangled with the parachute, the parachute ate platform. The cherts contain tridimensionally
could have covered the lander, thus preventing preserved filamentous and coccoidal (spheroidal)
it from opening. At the beginning of 2015 it was ▶ microfossils, including fossilized colonies of
announced that the orbital camera around Mars, microscopic pigmented cells. The distribution
HiRISE, had found the lander intact on the mar- and pattern of division of these later microfossils
tian surface, within the expected landing ellipse (called Eoentophysallis belcherensis) suggest
in Isidis Planitia but, unfortunately, it appeared to a relationship to the extant genera of
be only partially deployed. However, it can be ▶ cyanobacteria Entophysallis. These microfos-
demonstrated that the entry, descent and landing sils represent some of the oldest remains of
system did indeed work. identified cyanobacteria, together with cyst-like
cyanobacteria microfossils (akinetes) from
Gabon dated at 2.1 Ga (Fig. 1).
http://www.beagle2.com/index.htm
See Also
▶ Chert
▶ Cyanobacteria
Belcher Group, Microfossils
▶ Microbial Mats
▶ Microfossils
Palaeobiogeology-Palaeobotany-
▶ Sedimentary Rock
▶ Stromatolites
Definition
Benthic Mats
The Belcher Group comprises ▶ sedimentary
rocks from the Belcher Islands, Canada, dated at ▶ Microbial Mats
254 Benzene
▶ Planetary Nebula
Benzene ▶ Titan
William M. Irvine
Cernicharo J, Heras AM, Tielens AGGM, Pardo JR,

Synonyms Herpin F, Guélin M, Waters LBFM (2001) Infrared
space observatory’s discovery of C4H2, C6H2, and
C6H6; Cyclohexa-1,3,5-triene benzene in CRL 618. Astrophys J 546:L123–L126
Fouchet T, Bezard B, Encrenaz T (2005) The planets and
titan observed by ISO. Space Sci Rev 119:123–139
Woods PM, Millar TJ, Herbst E, Zijlstra AA (2003) The
Definition chemistry of protoplanetary nebulae. Astron
Benzene is an ▶ aromatic hydrocarbon in which
the six carbon atoms are arranged in a ring, with
all carbon bonds equal and intermediate in length
between single and double bonds. Under standard
laboratory conditions benzene is a colorless and Bernal’s Conception of Origins of Life
highly flammable liquid with a sweet smell. Ben-
zene is a known carcinogen and has various toxic Stéphane Tirard
effects on humans. Centre François Viète d’Histoire des Sciences et
des Techniques EA 1161, Faculté des Sciences et
des Techniques de Nantes, Nantes, France
History
Benzene was first isolated by the English chemist/ Keywords

physicist Michael Faraday in 1825, although the
natural aromatic resin that contains benzene comes Clays; Catalysis
from Southeast Asia and was known to Arab
traders during the Middle Ages. The cyclic struc-
ture of benzene was announced in 1865 by the History
German chemist Friedrich A. Kekulé, with the
data on carbon bond lengths coming from X-ray John Desmond Bernal was a pioneer of diffrac-
diffraction measurements. An astronomical detection X-ray method. His interest for biology
tion in the protoplanetary nebula CRL 618 has increased during the 1930s and the 1940s, prob-
been reported by Cernicharo et al. (2001) from ably in relation with the study of biological mol-
mid-infrared observations, and the measured abun- ecules (peptides, nucleic acids, etc.) with this new
dance is matched by chemical models (Woods physical method.
et al. 2003). In the Solar System benzene has His work on the origin of life was marked by
been observed in the atmospheres of Jupiter, Saturn his first publication on this topic in 1951: a little
and Saturn’s moon Titan (Fouchet et al. 2005). book entitled The Physical Basis of Life, which
came from a previous lecture (1947) and from a
paper published in 1949 in the Proceedings of the
See Also Physical Society.
In this text, Bernal gave a synthesis of previ-
▶ Aromatic Hydrocarbon ous theories, that is, Oparin’s (1924, 1938),
▶ Molecules in Space Haldane’s (1929), or Dauvillier’s (1947) ones.
Beta Pictoris b 255
Such as he claimed that primitive atmosphere of

earth contained CO2, he described the possibility Beta Pictoris b
of a progressive production of organic molecules
and finally of life. Daniel Rouan1 and Nader Haghighipour2
1
His main and original assumption regarded LESIA, Observatoire Paris-Site de Meudon, B
problems of dispersion and catalysis. Indeed, Meudon, France
2
Bernal claimed that fundamental reactions could Institute for Astronomy, University of Hawaii-
exist on clay deposits, marine, and freshwater, Manoa, Honolulu, Hawaii, HI, USA
which could insure confinement and catalysis.
During the 1950s and 1960s, he actively par-
ticipated in the scientific debates on the origin of
Synonyms
life and was very active in the diffusion of ideas
on the origin of life.
Bet Pic b
See Also
Definition
▶ Haldane’s Conception of Origins of Life
▶ Oparin’s Conception of Origins of Life Beta Pictoris b is a massive exoplanet directly
▶ Origin of Life detected around the A-star Beta Pic, on an orbit
with a semimajor axis between 8 and 10 astro-
nomical units. This is the smallest orbital distance
known so far among the small set of exoplanets
References and Further Reading observed by direct imaging. The star which is at a
Bernal JD (1951) The physical basis of life. Routledge and
distance of 20 pc (60 light years) is one of the
Kegan Paul, London best-known examples of a star surrounded by a
Bernal JD (1967) The origin of life. Weidenfeld and dusty ▶ debris disk. The disk was the first to be
Nicholson, London imaged and is now known to extend up to about
Dauvillier A (1947) Genèse, nature et évolution des
1,000 AU. The planet has a mass of about 9 Jupi-
planètes. Hermann, Paris
Haldane JBS (1929) The origin of life. The Rationalist ter masses and the right mass and location to
Annual, London, pp 242–249 explain the observed warp in the inner parts of
Oparia AI (1924) Proishkozhdeute Zhiui (The origin of the disk (Fig. 1).
life), Auu Syage Tvaus. In: Bernol JD (ed) The origin
A team led by A.M. Lagrange used the NAOS-
of Life. Weidenfeld and Nicolson, London, 1967
Operia AI (1938) The origin of life. MacMillan, CONICA instrument, an ▶ adaptive optics
New York corrected near-infrared imaging system mounted
on one of the 8.2-m Unit Telescopes of ESO’s
Very Large Telescope (▶ VLT), to observe Beta
Pic once in 2003 and then several times after
2008. On 2003 images, a faint source inside the
Bet Pic b disk was seen, but it was not considered as a
planet as it could have been a background star.
▶ Beta Pictoris b On the images taken in 2008 and spring 2009, the
source had disappeared, while in the observations
taken during autumn 2009 and later, the object
appeared on the other side of the disk after a time
Beta Electrons fully consistent with the object being an
exoplanet orbiting its host star. The size of the
▶ Beta Rays orbit was also consistent with this hypothesis.
256 Beta Rays
Beta Pictoris b,
Fig. 1 Beta pictoris
Since then, the spectral energy distribution in ▶ HR 8799: The First Directly Imaged Multi-
the infrared was obtained which showed the planet System
planetary parameters to be fully consistent with ▶ VLT
the first evaluation of the orbit and mass and
pointed to a dusty planetary atmosphere at
1,700 K.
Because the host star is young (12 million Beta Rays
years old), this discovery is considered as a strong
indication that ▶ gas giant planets can form Jun-Ichi Takahashi
within protoplanetary disks in only a few million NTT Microsystem Integration Laboratories,
years, a short time compared, for instance, to the Atsugi, Japan
age of the solar system (4.5 Gyr).
Keywords
See Also
Beta decay; Chirality; Radioactive particles;
▶ Adaptive Optics Weak interaction
▶ Debris Disk
▶ Direct-Imaging, Planets Synonyms
▶ Gas Giant Planet Beta electrons
Big Bang Nucleosynthesis 257
Definition From the standpoint of astrobiology, one of

the hypotheses for the origin of biomolecular
A beta ray is a stream of beta particles (electrons ▶ chirality, the so-called cosmic scenario, states
or positrons). In nuclear physics, beta decay is a that asymmetric energy sources in space-induced
type of radioactive decay in which a beta particle asymmetric chemical reactions of precursors in B
is emitted. interstellar dust, resulting in the ▶ enantiomeric
excess of terrestrial bioorganic compounds. It has
been proposed that beta rays are one of the can-
Overview didates for the source of such asymmetric chem-
ical reactions, which might lead to the origin of
In the case of electron emission, the decay is biomolecular chirality in terrestrial organic
referred to as beta minus (b), while in the case compounds.
of a positron emission as beta plus (b+). In beta
minus decay, a neutron is converted to a proton,
an electron, and an antineutrino; in beta plus
decay, a proton is converted to a neutron, a pos- See Also
itron, and a neutrino:
▶ Alpha Rays
n ! p þ e þ n e ▶ Asymmetric Reaction, Absolute
▶ Enantiomeric Excess
p ! n þ e þ þ ne ▶ Gamma Rays
▶ Homochirality
If the proton and neutron are part of an atomic
nucleus, these decay processes transmute one
chemical element into another. Beta decay does
not change the number of nucleons in the nucleus
but changes only its charge. For example:
BIF

90
Sr! Y þ e þ ne
90
▶ Banded Iron Formation
The kinetic energy of beta particles has a contin-
uous spectrum ranging from 0 to maximal avail-
able energy, which depends on parent and
daughter nuclear states participating in the
decay. A typical maximal available energy is Big Bang Nucleosynthesis
around 1 MeV, but it can range from a few keV
to a few tens of MeV. The most energetic beta Alain Coc
particles are ultra-relativistic, with speeds very Centre de Sciences Nucléaires et de Sciences
close to the speed of light. de la Matière (CSNSM) CNRS/IN2P3,
The spin of the electrons or positrons in beta Université Paris Sud 11, UMR 8609,
rays is longitudinally polarized due to parity Orsay, France
nonconservation in the weak interaction medi-
ated by charged W particles. The helicity of a
beta electron, that is, the spin angular momentum Keywords
component of the kinetic momentum direction, is
negative (left-handed), and that of a beta positron Big bang, Deuterium, Helium; Lithium;
is positive (right-handed). Nucleosynthesis
258 Big Bang Nucleosynthesis
Synonyms and a tiny fraction of neutrons and protons.

Weak reactions (with electrons and neutrinos)
Primordial nucleosynthesis maintained equilibrium between the number of
protons and neutrons until the Universe cooled
down to 1010 K. This occurred because the rate of
Definition weak reactions became slower than the rate of
space expansion. Consequently, the ratio of the
The nucleosynthetic process that took place number of neutrons to protons became frozen.
within the first 20 min after the Big Bang is called When the temperature dropped to 109 K, the
Big Bang Nucleosynthesis (BBN) or Primordial fusion of a proton and a neutron leading to a
Nucleosynthesis. At this early epoch, the Uni- deuterium nucleus became favored compared to
verse was dense and hot enough to allow for deuterium dissociation by high-energy photons.
nuclear reactions to take place, producing the This was the starting point of primordial nucleo-
“light elements”: 4He, 2H (D, i.e., deuterium), synthesis that stopped, after 20 min, with the
3
He, and 7Li, starting from neutrons and protons. formation of 7Li because no nucleus with mass
The comparison between the primordial abun- 8 (or 5) exists and because of the decreased den-
dances of these isotopes, deduced on one hand sity and temperature. Only the 4He, D, 3He, and
7
from observations and on the other hand from Li isotopes are produced in Standard Big Bang
model calculations, is one of the main supports Nucleosynthesis, involving a dozen main nuclear
of the Big Bang model. reactions.
The comparison between the calculated Big
Bang Nucleosynthesis isotopic abundances and
History those deduced from observations in primitive
astrophysical sites was used to determine the
Prominent landmarks in the development of the density of ordinary matter in the Universe. It is
Big Bang Nucleosynthesis theory include works now more precisely deduced from the observa-
by Gamow in the 1940s (out of equilibrium tions of the anisotropies of the cosmic back-
nucleosynthesis in an expanding Universe domi- ground radiation. Despite the fact that the
nated by radiation), Peebles in 1966 (Big Bang primordial abundances of these light isotopes
Nucleosynthesis calculations up to 4He), and span nine orders of magnitude, the agreement
Wagoner in 1973 (Big Bang Nucleosynthesis between calculations and observations is good
calculations including 7Li). with the, yet unexplained, exception of 7Li (but
within a factor of 3–5). According to models,
the Big Bang Nucleosynthesis contributions to
Overview the solar system abundances amount to 90 %
for 4He and 25 % for 7Li and are uncertain for
3
The Big Bang model is supported by three pieces He but are remarkably the only source of
of observational evidence: the expansion of the present-day deuterium.
Universe, the ▶ cosmic background radiation, All the parameters of the Standard Big Bang
and the Primordial or Big Bang Nucleosynthesis. Nucleosynthesis are now known from other
In the framework of an expanding Universe, with sources. It is now used as a probe of nonstandard
uniform temperature and density that decrease physics in the early Universe.
with time, a temperature of 1011 K is reached a
fraction of a second after the Big Bang.
According to our present knowledge, the only See Also
particles present in the Universe at this time
were photons, electrons, positrons, neutrinos, ▶ Cosmic Background Radiation
and antineutrinos, all in equivalent numbers, ▶ Nucleosynthesis, Explosive
Binary Stars, Young 259
▶ Nucleosynthesis, Neutrino
▶ Nucleosynthesis, Stellar Binary Stars, Young
Steven W. Stahler
Department of Astronomy, University of B
Coc A, Goriely S, Xu Y, Saimpert M, Vangioni E (2012) California, Berkeley, CA, USA
Standard big bang nucleosynthesis up to CNO with an
improved extended nuclear network. Astrophys
J 744(1–18):158 Keywords
Fields BD (2011) The primordial lithium problem. Annu
Rev Nucl Part Sci 61:47–68
Iocco F, Mangano G, Miele G, Pisanti O, Serpico PD Star formation
(2009) Primordial nucleosynthesis: from precision
cosmology to fundamental physics. Phys Rep 472:1–76
Peebles PJE (2009) Finding the big bang. Cambridge
University Press, Cambridge. ISBN 13:978- Definition
0521519823
Weinberg S (2008) Cosmology. Oxford University Press, Most stars are not isolated objects, but have an
Oxford. ISBN 13: 978-0198526827 orbiting companion. This basic fact holds not only
for mature stars but also for objects at an earlier
stage of evolution. Indeed, several young star clus-
Bimolecular Reaction ters have a higher fraction of binaries than do main-
sequence stars generally. Young binaries exhibit a
Steven B. Charnley very broad range in separations and therefore
Solar System Exploration Division, Code 691, orbital periods. In the widest pairs, the orbits are
Astrochemistry Laboratory, NASA Goddard highly eccentric. Conversely, the tightest pairs are
Space Flight Center, Greenbelt, MD, USA locked into circular orbits. Planet-forming disks are
absent in these latter systems, apparently because of
the disturbing influence of the companion.
Synonyms
Two-body reaction Overview
Astronomers have long known that most stars

Definition have binary partners, that is, companions locked
gravitationally into orbit. About 60 % of solar-
A bimolecular reaction is a chemical process type, main-sequence stars have at least one such
involving two reactants. Reactants may be elec- companion. The vast majority of these multiple
trons or atoms and molecules existing in various systems are binaries, but triples and even quadru-
combinations of charge states (neutral, anionic, ples also exist. Binarity is also common for stars
or cationic). The most important reactant combi- of other spectral types.
nations for interstellar chemistry are ion-neutral, Some binary companions are so close together
neutral-neutral, anion-neutral, electron-neutral, that one can detect the induced wobble in the
and electron-ion. stars’ motion, through a periodic Doppler shift
in the wavelength of spectral lines. Such spectro-
scopic binaries are relatively rare. Most systems
See Also are discovered because the two stars share a com-
mon spatial motion. This implies that their
▶ Anion pairing is physical and not a chance superposi-
▶ Interstellar Chemical Processes tion. Overall, the observed range of periods is
260 Binding Constant
vast, from less than a day to millions of years, Ghez AM, Neugebauer G, Matthews K (1993) The multi-
corresponding to separations from 0.01 to plicity of T Tauri stars in the star-forming regions
Taurus-Auriga and Ophiuchus-scorpius: A2, 2 micron
10,000 AU. speckle imaging survey. Astron J 106:2005
The fact that most mature stars have compan- Jensen ELN, Mathieu RD, Fuller GF (1994) A connection
ions naturally leads us to wonder if this situation between submillimeter continuum flux and separation
held further back in time. Are ▶ pre-main- in young binaries. Astrophys J 429:L29
Zinnecker H, Mathieu RD (eds) (2001) The formation of
sequence stars, those too young to fuse hydrogen binary stars. Astronomical Society of the Pacific, San
into helium, also preferentially found in binaries? Francisco
What about even younger objects, those still
gathering mass from their parent molecular
clouds? Since the 1980s, many researchers have
investigated the matter, and the answer is now
clear. Pre-main-sequence stars are also very Binding Constant
likely to have a binary companion. Indeed, the
binary fraction in some young clusters is even ▶ Affinity Constant
greater than for main-sequence stars in the field.
The systems being found show a wide range in
orbital separations. Some are tight enough to be
detected as spectroscopic binaries. Most young Binding Energy
binaries, however, are noticed because they
exhibit a common spatial velocity within their Steven B. Charnley
parent stellar group. On the whole, pre-main- Solar System Exploration Division, Code 691,
sequence binaries span a wide range of separations Astrochemistry Laboratory, NASA Goddard
and periods, just as do main-sequence systems. Space Flight Center, Greenbelt, MD, USA
Many individual pre-main-sequence stars
have planet-forming disks, as evidenced by their
excess infrared and millimeter emission. The Definition
same is true for stars within binaries, but with a
significant caveat: if the binary separation is less Binding energy is the energy required to disas-
than about 100 AU, corresponding to a period of semble an entity into its constituent parts. This
about 1,000 years, then the excess emission is corresponds to the mechanical work which must
absent. Apparently, the relatively nearby com- be done in acting against the forces which hold
panion star prevents formation of any circumstel- the entity together. The binding energy of an
lar disk. For periods as low as a few days, the atom is that required to disassemble an atom
primary and its companion are locked into per- into free electrons and a nucleus, acting against
fectly circular orbits. the electromagnetic force. The nuclear binding
energy is that required to disassemble a nucleus
into its constituent neutrons and protons, acting
See Also against the strong nuclear force. The term is also
used in other contexts, for example, for the
▶ Pre-Main-Sequence Star energy involved in attaching a gaseous molecule
to an ▶ interstellar dust grain upon collision. In
this case, the magnitude depends on both the
References and Further Reading physical nature of the molecule (e.g., polarizabil-
ity) and of the grain surface. Binding energies for
Duquennoy A, Mayor M (1991) Multiplicity among solar-
▶ physisorption (through van der Waals bond-
type stars in the solar neighborhood II: distribution of
orbital elements in an unbiased sample. Astron ing) are generally much lower than those of
Astrophys 248:485 ▶ chemisorption.
Biobarrier 261
See Also international level; and the establishment of col-

laborative programs with other international sci-
▶ Chemisorption entific societies with related interests.
▶ Interstellar Dust In the International Astronomical Union, this
▶ Physisorption Commission is part of Division F, which was B
renamed in 2012 to be Planetary Systems and
Bioastronomy.
Bioastronomy
History
▶ Astrobiology
The International Astronomical Union’s Commis-
sion 51 was established in 1982 as “Bioastronomy:
Search for Extraterrestrial Life”, was renamed
Bioastronomy (IAU Commission 51) simply “Bioastronomy” in 2006, and renamed
again “Astrobiology” in 2015. From an early con-
William M. Irvine centration on ▶ SETI, bioastronomy has expanded
University of Massachusetts, Amherst, MA, USA its interests as described above.
Definition
See Also
In current usage, bioastronomy is both a synonym
for astrobiology (although the term was intro- ▶ IAU
duced before NASA coined “astrobiology”) and ▶ SETI
the former title of Commission 51 of the Interna-
tional Astronomical Union (▶ IAU). Commis-
sion 51, which has recently been renamed
Astrobiology and which may be re-numbered Biobarrier
following the IAU General Assembly in August,
2015, defines its field to be the study of the origin, Catharine A. Conley
evolution, and distribution of life in the universe. NASA Headquarters, Washington, DC, USA
In this context, bioastronomy/astrobiology
encompasses the search for extant life, evidence
of past life, or evidence of prebiotic chemistry on Definition
solar system bodies, including Mars, Europa,
Titan, and Enceladus; the search for planets In ▶ planetary protection, a biobarrier is a
around other stars and potential spectroscopic mechanical barrier to protect a spacecraft or asso-
evidence for habitability and biological activity; ciated component(s) against microbial
the origin of the biogenic chemical elements and recontamination following the application of
the study of biologically relevant molecules in ▶ bioburden reduction procedures.
the interstellar medium and in primitive solar
system objects such as comets, undifferentiated
asteroids, and some meteorites; the search for See Also
intelligent signals of extraterrestrial origin
(▶ SETI); the study of the origin, early evolution, ▶ Bioburden
and environmental constraints for life on Earth; ▶ Bioburden Reduction
the coordination of efforts in all these areas at the ▶ Planetary Protection
262 Biobloc
microbial reduction (▶ DHMR), ultraviolet or

Biobloc g-ray irradiation, and treatment with a
sterilizing gas.
▶ Biostack
See Also
Bioburden
▶ DHMR
Catharine A. Conley ▶ Pasteurization
NASA Headquarters, Washington, DC, USA ▶ Planetary Protection
▶ Sterilization
Definition
In ▶ planetary protection, the bioburden is the Bioburden-Controlled Environment

total amount of viable microorganisms sitting
on the surface and inside a spacecraft. The Catharine A. Conley
bioburden is evaluated using ▶ assays in or on NASA Headquarters, Washington, DC, USA
items of interest. To evaluate or measure the
bioburden, the assays can be standardized by
specific procedures. Definition
In ▶ planetary protection, a bioburden-controlled

environment is a place where the number of via-
See Also ble ▶ microorganisms, and therefore the poten-
tial for contaminating spaceflight hardware, is
▶ Assay controlled and minimized. ▶ Clean rooms, lami-
▶ Microorganism nar flow hoods or cabinets, and other environ-
▶ Planetary Protection ments in which the quantity of microorganisms
and/or particulates is monitored and maintained
at a specified level are considered “bioburden
controlled.”
Bioburden Reduction
Catharine A. Conley
See Also
▶ Bioburden
▶ Clean Room
Definition
▶ Planetary Protection
Bioburden reduction involves any activities
designed to remove or destroy ▶ microorganisms
that are performed in order to reduce ▶ bioburden
levels on or in an item of interest. These activities Biodetection System
could involve cleaning and wiping with appropri-
ate alcohol or chemical solutions, dry heat ▶ Biosensor
Biodiversity 263
quality and variety of living organisms in relation

Biodiversity with the threats constituted by ▶ environment
degradation, species extinction, and potential
Simon Tillier and Guillaume Lecointre losses of socioeconomic benefits (Wilson 1988).
Département Systématique et Evolution, UMR In parallel, the convergence of interest from the B
7138 CNRS-MNHN-UPMC-IRD, Muséum scientific community, from the nature conserva-
National d’Histoire Naturelle, Paris, France tion organizations, and from the southern coun-
tries, who want to control the economic benefit
from their living resources, led the United
Keywords Nations Environment Programme (UNEP) to
explore the need for an international convention
Ecology; Ecosystem services; Environment; on biological diversity. This approach led to an
Evolution; Systematics; Taxonomy international legal instrument for the conserva-
tion and sustainable use of biological diversity:
the Convention on Biological Diversity was
Synonyms opened for signature on June 5, 1992, at the
United Nations Conference on Environment and
Diversity of life Development, better known as the Rio “Earth
Summit,” and was adopted by 168 countries
within the next year. In terms of scientific orga-
Definition nization and programming, an international
research program on biodiversity, named
Biodiversity designates the condition of life on Diversitas, was established in 1991 jointly by
Earth in terms of its variation at all levels of UNESCO, SCOPE (Scientific Committee on
biological organization, from genes to ▶ ecosys- Problems of the Environment or the International
tems. By extension, it is used to designate life on Council of Scientific Unions), and the IUBS
Earth itself, most generally at the ▶ species level (International Union of Biological Sciences) to
but often also at all organization levels altogether. “promote an integrative biodiversity science,
The political impact of the term led to a definition linking biological, ecological and social disci-
adopted by the International Convention on Bio- plines in an effort to produce socially relevant
logical Diversity (Art.2): “Biological diversity’ new knowledge; and provide the scientific basis
means the variability among living organisms for the conservation and sustainable use of biodi-
from all sources including, inter alia, terrestrial, versity.” Ten years after Rio, the second Earth
marine and other aquatic ecosystems and the Summit held in Johannesburg in 2002 set the
ecological complexes of which they are part; so-called 2010 target: “to achieve by 2010 a
this includes diversity within species, between significant reduction of the current rate of biodi-
species and of ecosystems” (Convention on Bio- versity loss at the global, regional and national
logical Diversity 1992). level as a contribution to poverty alleviation and
to the benefit of all life on Earth.” Clearly the
target has not been met, in the absence of instru-
History ments of measure as well as of effective policies.
However, the conjunction of scientific interest
Although first introduced in the end of the 1960s, and social pressures has modified how ecologists,
the word biodiversity, as a contraction of biolog- taxonomists, population geneticists, paleontolo-
ical diversity, has been coined by Walter gists, and social scientists of environment situate
G. Rosen while preparing a national US scientific their own disciplines, toward an integrated vision
forum held in Washington in 1986. The objective where most see themselves as contributors to
of this forum was to discuss the importance of the biodiversity research.
264 Biodiversity
Overview not catastrophic at the scale of the human

lifespan, but rather in a time span of several
Biodiversity science is above all integrative. The millennia, which we do not perceive directly but
central problem of biodiversity research is to is still a very short instant in the more than 3.5
understand how diversity of life, that is, the var- billion year-long history of life on Earth. This
iation in quality and quantities of living things at crisis, sometimes referred to as the Sixth Extinc-
any scale, relates to the functioning and evolution tion, is likely the direct result of human activities.
of ecosystems including the human populations Understanding the patterns and processes of
and societies. In consequence, all population- biodiversity loss and change is crucial for our
level research may be relevant at any scale, species, not only because of the aesthetic, ethical,
from any local group of living organisms to all or cultural values attached to biodiversity but also
past, present, and future life. Taxonomists because it could have numerous far-reaching con-
(or systematists, including paleontologists) sequences for our own life-support system. Even
explore and describe the patterns of biodiversity, the chemical composition of the air we breathe
present and past, proposing scenarios illustrating and of seawater depends upon biological activity
“how” – in terms of a suite of inferred historical well beyond the increase of carbon dioxide in the
events – biodiversity reached its present state at atmosphere and of acidity in the seawater, which
the largest scales. Evolutionary biologists aim at results directly from human activities.
understanding “how” – in terms of biological One of the likely consequences of
processes – biodiversity reached its present state man-induced changes in the environment is the
at genetic and population levels. Ecologists reduction in the capacity of natural and managed
explore the patterns of species interactions in ecosystems to deliver ecological services, such as
ecosystems and the mechanisms of ecosystem the production of food and fiber, carbon storage,
functioning; and social scientists explore the rela- nutrient cycling and resistance to climate, and
tionships between human populations and socie- other environmental changes. Assessing the
ties and nonhuman biodiversity. Focus on causes and consequences of biodiversity changes
socioeconomic benefits, which were already and establishing the bases for the conservation
encompassed by the initial thoughts in the and sustainable use of biodiversity are major
1980s, has brought emphasis of the concept of scientific challenges of our time. The major ques-
ecosystem services, that is, the part of the ecosys- tions that research is facing are summarized in the
tem functions and products that is used by our Diversitas research program (Diversitas 2002,
species. 2010).
We understood as late as in the 1980s that the
number of living species is probably tenfold the How Did Biodiversity Evolve in Space and
previous estimates, in the 10–40 million living Time to Reach the Current State?
species range, rather than 1–2 million (Erwin There are two kinds of sciences answering the
1982 and subsequent papers). In parallel, it has “how” questions. The first one is focusing on the
become progressively evident that unprecedented biological processes by which living matter
losses and changes in biological diversity are diversifies its forms (see the entry Evolution
taking place at the genetic, species, and ecosys- (biological)). The way these scientists demon-
tem levels, particularly in nonmarine groups of strate their statements is close to that of chemistry
organisms so far. Comparison with major extinc- or physics. By manipulating populations of
tion crises in the geological history of the Earth organisms with very short generation times, it is
reveals that the current rate of extinction is at possible to develop a hypothetico-deductive
least in the same order of magnitude and several experimental approach to these “how” questions:
hundred times greater than the average between population genetics is of first importance in the
crises (Dirzo and Raven 2003). The present field. But there are also other experimental sci-
extinction crisis does not strike us because it is ences like causal embryology or physiology that
Biodiversity 265
participate to the understanding of this “how,” very few species interact. The historical “how”
though without any population approach. may be easier to grasp, thanks to both paleonto-
The second approach is the historical “how.” logical observations and, possibly, by combining
A set of sciences like descriptive and compara- the phylogeny of the species composing a com-
tive embryology and anatomy, paleontology, and munity with the composition of the latter through B
systematics participate in a historical interpreta- time. An objective is to relate ecosystem change
tion of how life arrived to its present state. The with causal factors such as climate change, entry
present distribution of organisms is explained by of a clade into a new region, or the origin of
a suite or series of events, not necessarily linked innovations in a clade.
to each other by causal relations. ▶ Phylogeny
provides part of the information needed for the How Much Biodiversity Exists and How Does
ordering of events through time. Other data, pale- Its Change or Loss Affect the System
ontological, stratigraphical, geological, etc., also as a Whole?
participate in our knowledge of the history of A prerequisite to the “how” is the “what,” that is,
biodiversity. The answer to the historical “how” documenting biodiversity. Understanding how
has made decisive progress in the last two little we know about the existing species implies
decades through the association of the phyloge- that traditional methods and techniques are inap-
netic method introduced by Willi Hennig propriate, and new strategies are developed to
(Hennig 1966) with informatics and use of DNA increase the efficiency of field sampling, the qual-
sequences: phyloinformatics now allows ity and accessibility of collections for fieldwork,
processing large data sets, whether molecular or the collection and curation of voucher specimens,
morpho-anatomical. Thanks to phylogenetic the technologies for imaging and DNA sequenc-
methods applied to molecular characters, our ing, the digitization of legacy data, and the devel-
vision of the tree of life has been radically mod- opment, maintenance, and connectivity of
ified within the last 25 years and is still evolving. relevant databases. The need for rapid-capture
Within 25 years, our vision of the tree of life has technologies for identifying known species and
shifted from five “kingdoms” to three main discovering new ones has led to the development
branches or clades, of which two are bacteria of the “Barcode of Life” initiative, which aims at
domains diverging from each other just as much providing IT tools for identifying species through
as from ours, the eukaryotes (the organisms con- short DNA sequences obtained from specimens
stituted by one to many cells with a nucleus). identified by an expert and kept in legacy collec-
Convincing evidence has been brought to explain tions (Consortium Barcode of Life 2010; Golding
the origin of eukaryotes by multiple bacterial et al. 2009). The need for access to the informa-
endosymbioses: our own cells, their mitochon- tion on species occurrences and on the collec-
dria, and plasmids originate indeed from as tions, which may well soon include the only
many bacterial lineages. It is now admitted that remaining specimens of many species in view of
lateral ▶ gene transfer (i.e., gene transmission the current rate of extinctions, has led to the
between two otherwise independent lineages) is constitution of the so-called GBIF (Global Bio-
a relatively frequent phenomenon at least in bac- diversity Information Facility 2010; Edwards
teria and may well influence significantly their et al. 2000), which is an international informatics
evolution. infrastructure that allows the interoperability of
At the ecosystem level, “how” is also a ques- the innumerable databases on collection speci-
tion from both points of view: from an ecological mens and species occurrences worldwide.
processes point of view, not only chance, that is, A serious problem for assessing species diver-
the initial conditions, but also species interactions sity is the uneven distribution of expertise among
doubtless shape the local communities in time, taxa and countries: groups of very high diversity
even though we are hardly able to understand and and ecological importance, such as nematode
even less to predict the change when more than worms, acarians, most insects, or microbes, in
266 Biodiversity
general, are notoriously understudied and very projected on a map to delimit species’ potential
poorly known, whereas groups that are closer to distribution in space. It is then easy to modify
us in terms of body organization, size, and appeal some environmental parameters, including cli-
to imagination may even have more specialists matic ones, to predict the consequences of
than species, as is the case of birds or elephants. changes on species occurrences and, to some
In parallel, expertise is located mainly in Western extent, on the composition of communities. This
developed countries, principally in Europe and approach has proved very powerful to define pol-
North America, whereas most megadiverse trop- icies and strategies regarding invasive pest spe-
ical countries have very few specialists and insuf- cies in Mexico or the design of protected areas;
ficient research infrastructures: biodiversity is in but it is somewhat rudimentary by not taking into
the South but expertise and infrastructures are in account ecological structures and processes, nor
the North. metapopulation dynamics, habitat fragmentation,
For their own biodiversity assessments and in etc. The ultimate goal is not just to understand but
response to the environmental concerns, which are to predict biodiversity change, developing biodi-
at the origin of environmental regulations, rich versity scenarios that predict biodiversity change
countries have developed observation networks at the landscape, regional, and global scales in
that are feeding large observational databases, response to various scenarios of how anthropo-
managing billions of observational data. These genic drivers will change in the future.
databases provide the basic tool for the application
of environmental regulations regarding species How Does Biodiversity Correspond to the
and habitats but also constitute a tool for research Delivery of Ecosystem Functions and Services
when standardized and made interoperable, inter and What Is the True Value of These
alia, thanks to the GBIF. Their weakness is that Commodities?
they necessarily concentrate on taxa for which Over two decades of research in ecology have
there are a number of observers, that is, principally shown that the combination of abiotic (physical
higher plants and vertebrates. However, when and chemical) factors together with biological
enough data have been collected in the medium interactions determines the limit of biodiversity
and long term, it has been possible to show such in a community and that the composition of this
phenomena as the decline of populations of com- community influences, in turn, the way it func-
mon birds in Europe or the shift in distribution tions. A large number of experiments since the
toward the north of bird and plant species as a beginning of the 1990s have confirmed that gen-
consequence of global warming. In the domain erally the increase in biodiversity has a positive
of fisheries, fishing data collected since the end effect on ecosystem functioning.
of the nineteenth century have allowed us to show Three mechanisms have been proposed to
that overfishing started at least more than one explain this positive effect. The first is a simple
century ago, leading to constant diminution of sampling effect: species-rich ecosystems would
tonnage and size of the fish at the world level and produce more biomass, simply because the prob-
unpredictable consequences over the food web ability for including a highly productive species
and equilibrium of the global ocean. is higher as the number of species is increased.
However, assessing is not enough in both sci- The second is the functional complementarity of
entific and societal terms: we want to be able to species resulting from a favorable pattern of traits
predict the change, for scientific reasons as well in the community: by increasing diversity, one
as for decision making in environmental policy. increases the number of effective ecological
The solution lies in modeling, operationally functions as more ecological niches are
based so far upon statistics on species distribution exploited. For example, two plant species having
in terms of environmental parameters. This different root lengths extending down to the
approach, called niche modeling, allows one to ground will better exploit resources in the soil
calculate an ecological envelope that can be and then produce more biomass together, than a
Biodiversity 267
single of them. Finally, interaction of two spe- is also suspected that diversity favors diversity.
cies, then called facilitation, may have a positive An illustration of this is, for example, the recent
effect on production: for example, brambles in a discovery of very high levels of horizontal or
meadow may create favorable conditions for lateral transfers of DNA in microbial communi-
smaller species that need moisture and shade, ties, in a way that genomes exhibit functional- B
such as arum lilies, and doing so favors the environmental patterns as well as historical pat-
increase in productivity of the system. terns. Horizontal heritage is far more important
These mechanisms may be effective at any than previously thought, compared to “classi-
time and space scale, from bacteria and fungi in cal” vertical heritage. These networks of perma-
a piece of cheese to the entire tropical forests and nent DNA exchanges among a diversity of
oceans. However, for practical reasons, most microbial “species” in a given environment
experiments have been based on plant communi- (either marine, lacustrine, or terrestrial) stabilize
ties, including the two best known, the European the role played by these environmental commu-
BIODEPTH and the American Cedar Creek nities in the biosphere. Generally speaking, the
experiments, which both have shown a positive question of positive feedback of diversity is fun-
effect of diversity upon biomass accumulation in damental in view of the present changes in eco-
the communities. systems and still requires further research at
The relationship between diversity and stabil- various levels of ecosystem organization. If it
ity of ecological communities has also been stud- is true, human activities endanger self-
ied, showing at least theoretically that some regulation of ecosystems, of which many are at
species may have a key role for the persistence risk of collapsing with unavoidably serious risk
of the community when facing environmental for our own species.
disruption: roughly, having more species
increases the chance of having at least one or a How Can Scientific Investigation Support
few able to develop under modified conditions Policy and Decision Making to Encourage
and in so doing allow persistence of the commu- More Sustainable Use of Biodiversity?
nity. Few experiments have tested this conclu- In view of the societal needs for a better under-
sion, but these few confirm so far the theoretical standing of the functions of biodiversity for
prediction of a relation between diversity and human beings (and well-being where possible),
stability of ecosystem functioning. communicating scientific results to relevant polit-
However, most experiments neglect the tro- ical levels is crucial. Researchers are trained in
phic relationships between species, which we research and communication with other
know are very complex in species-rich ecosys- researchers, but not in communication toward
tems, from predation and parasitism to mutual decision makers, which results in numerous mis-
dependence such as in pollination processes. understandings. This renders the formation of an
A starting point may be the top-down effect, ad hoc mechanism a necessity. Based on the
through which a predator controls the density of example of the IPCC (Intergovernmental Panel
prey, which themselves control production at on Climate Change), the United Nations and the
lower trophic levels. Diminution in density at G8 have endorsed in 2010 the creation of such a
the top levels may allow an indirect increase in mechanism, called the Intergovernmental
production at the lowest levels; but it may also Science-Policy Platform on Biodiversity and
have just the opposite effect when top levels Ecosystem Services (IPBES 2010; Larigauderie
participate significantly in nutrient recycling, and Mooney 2010) as initially proposed by
which itself increases productivity: this may France in 2005. The IPBES progresses at an
well be what happens as mankind clearly exploits intergovernmental pace, which is slower than
and overexploits large predators. species extinctions and ecosystem change, but
Overall, we know now that diversity favors will hopefully allow progress in the implementa-
ecosystem resilience and biomass production. It tion of the necessary policies.
268 Biodiversity
Key Research Findings systematics and evolutionary biology as central

disciplines along with inventories and ecological
Since the beginning of the 1980s, a few points modeling.
have been understood: Systematics is important in understanding the
reasons why we classify living and fossil organ-
isms as we do. Systematics provides the reason
• First is a revolution in our vision of the tree of
why the platypus is classified well apart among
life and of the origin and diversification of
mammals. Phylogenetic systematics explains
eukaryotes. All eukaryotes are no more nor
why the collection of traits exhibited by the platy-
less than each of the two main “prokaryotic”
pus is so unique. If a decision has to be made
clades (Bacteria and Archaea), fungi and ani-
concerning the preservation of the platypus, the
mals now appear as originating from a common
ecological role of the platypus is going to have a
ancestor not shared with other groups, and
negligible weight compared to the argument from
humans are nothing more than the tip of a tiny
systematics. The platypus (or the coelacanth,
twig among all mammals, which occupy noth-
choose the one you prefer) is negligible in terms
ing more than a small branch of the small clade
of biomass, and its role in its environment could
constituted by all animals in the tree of life.
be replaced by another imported species. In terms
• Next is the change of paradigm in both taxon-
of what organisms do in their environment, the
omy and ecology: we now know that we know
platypus (or the coelacanth) can perfectly well be
nothing about most living species and that the
sacrificed. By contrast, phylogenetic systematics
way we interact with biodiversity is not sus-
will provide the arguments to regard this species
tainable for the present ecosystems at any
as a very important one. Phylogeny will show that
level, whether marine or land based.
the platypus is the product of a very ancient
• The whole biosphere is changing at all levels,
mammal lineage represented today by three spe-
possibly faster than ever, and surely through a
cies only. The characters exhibited by the platy-
process without precedent, which is provoked
pus show this: it lays eggs and it has a bizarre
by our species: destruction of whole ecosys-
anatomy of the temporal area. In parallel, the
tems and modification of others through
lineage gained characters on its own branch,
anthropic activities including transport of spe-
such as a duck-like beak or venomous fangs on
cies are realities that we have to face.
the posterior limbs. The combination of the
• In scientific terms, we understand better how
whole set of properties makes the platypus a
biodiversity acts in the functioning of the whole
kind of unique evolutionary heritage – a rare
ecosystems: biodiversity favors both produc-
combination of characters. Protecting species
tion and stability of ecosystems and likely has
should therefore be considered in terms of what
a positive feedback on diversity itself.
they have (systematics), not only through what
they do (ecology).
Future Directions Evolutionary biology is also important
because we cannot pretend to understand and
On the purely scientific point of view, the main properly protect biodiversity without having a
future direction to maintain our understanding of deep knowledge of the processes that generate
biodiversity is to keep all contributing sciences biological diversity at all levels. This is going to
visible at the political level. The study of biodi- be important because it is not sufficient to save
versity is not merely a matter of counting the the present state of nature as if it would not be
species that are found in a given area or proposing going to change again. Life is continuously
models predicting responses of ecosystems to changing. It is important to protect the potential
anthropic perturbations. The future in studies ability to get adapted to new conditions, so
and understanding of biodiversity must keep maintaining genetic variability and our
Biodiversity 269
knowledge about processes that generate it (and • Complementary and necessary for develop-
select it afterward) is of key importance. ment of modeling is documenting what occurs
The second relevant direction for our knowl- and what happens, because without reliable
edge and action is to consider human societies data we may hardly hope to understand what
and their products as parts of biodiversity. There is happening globally. Even local predictions B
is a philosophical tradition that keeps humankind require global data sets because, like in mete-
separated from an undefined nonhuman “nature” orology, operational modeling is based upon
or an undefined category of “animal” and that statistical approaches of which precision and
keeps culture separated from nature. In contrast, reliability are in proportion to the amount of
we are not only trying to change the world to save observations available. This need for data and
the diversity of this wonderful life but also to save monitoring is taken into account by the
the cultural-material heritage for the few next projected construction of the Global Earth
human generations. Observation Biodiversity Observation Net-
Although the economics of biodiversity is a work (GEO BON), which is a program for
rapidly expanding field, and understanding the the coordination of observations on biodiver-
relationship between biodiversity and ecosys- sity promoted by Diversitas. GEO BON is a
tem functioning is still seen as a research prior- component of the Global Earth Observation
ity, for practical reasons focus is presently made System of Systems (GEOSS), an initiative
on observations and monitoring. Not only scien- led by the World Meteorological Organization
tists but also all kinds of decision and policy (Scholes et al. 2008).
makers need reliable information on what is
happening and projections of what may happen, Clearly, biodiversity research is shifting from
both locally and globally. Decreases of coral traditional fundamental questions on what is
reefs and more generally ecological conse- there, why it is there, and how it works to ques-
quences of acidification of the oceans, changes tions relating directly to societal concerns guided
in the composition of the atmosphere, invasion by the central question of where mankind is going
of exotic species, changes in the species distri- in a context of radical ecological change at the
bution resulting from climate warming and global scale. So far, this shift is beneficial rather
transportation by humans, and destruction of than detrimental to fundamental research; and
habitats under anthropogenic pressures are all indeed the questions raised by society regarding
happening and hardly evaluated due to the lack its own future as a constituent of biodiversity will
of appropriate instruments. This observation not find their answers if progress is not going on
calls for at least two lines for action from the in fundamental ecology, systematics, and evolu-
scientific community: tionary biology.
To conclude on a positive note, we are right
• The first one is improving our capacity for the to be concerned by the ongoing Sixth Extinction.
analysis and prediction of the change in biodi- However, life has to be considered in the very
versity. Operational models based upon niche long term. The Sixth Extinction is a problem of
modeling that are already in use to assist our responsibility for the next 2 to some
decisions should be developed further. The pre- 200 human generations, but it is not a problem
dictive value of models could doubtless still be for life itself. Several drastic extinctions have
improved by incorporation of ecosystem taken place in the past million years; there will
functional parameters, as ambitioned by many be others. Our theoretical framework allows us
ecologists; however, this development is to predict that biodiversity will be recovered, as
impeded so far by the weakness of the concepts rich as ever or more, and life and Earth will
relating to ecosystem functions, which still continue to change together as time will con-
requires a qualitative jump forward. tinue going by.
270 Biodiversity (Planetary Protection)
See Also
Biodiversity (Planetary Protection)
▶ Adaptation
▶ Domain (Taxonomy) Catharine A. Conley
▶ Ecosystem NASA Headquarters, Washington, DC, USA
▶ Environment
▶ Gene Definition
▶ Genome
▶ Lateral Gene Transfer For ▶ planetary protection, the biological diver-
▶ Phylogenetic Tree sity (biodiversity) in an environment is the inven-
▶ Phylogeny tory of types of ▶ microorganisms identified as
▶ Species being present. For microorganisms, the identifi-
cation depends also on the ▶ assay which is used.
For instance, to prepare future missions to
▶ Mars, the space agencies are conducting stud-
References and Further Reading ies in the clean rooms to be used for instrument
and spacecraft assembly.
Convention on Biological Diversity (1992) N 30619.
United Nations, treaty series, 1760, 1993. pp 142–308
Consortium Barcode of Life (2010) http://www.
barcodinglife.org/ See Also
Dirzo R, Raven PH (2003) Global state of biodiversity and
loss. Annu Rev Environ Resour 28:137–167
Diversitas (2002) Diversitas science plan. Diversitas, Paris ▶ Bioburden
Diversitas (2010) An international programme of biodiver- ▶ Bioburden Reduction
sity science. http://www.diversitas-international.org ▶ Planetary Protection
Edwards JL, Lane MA, Nielsen ES (2000) Interoperability
of biodiversity databases: biodiversity information on
every desktop. Science 289(5488):2312–2314
Erwin TL (1982) Tropical forests: their richness in Cole-
optera and other arthropod species, 3Coleopterists’. Bioenergetics
Bulletin 36:74–75
GBIF (2010) Global Biodiversity Information Facility.
http://www.gbif.org. Ricardo Amils
Golding GB, Hanner R, Hebert P (eds) (2009) Special Departamento de Biologia Molecular,
issue on barcoding. Mol Ecol Resour 9(supplement) Universidad Autónoma de Madrid, Madrid,
1:1–267
Spain
Hennig W (1966) Phylogenetic systematics. University of
Illinois Press, Urbana
IPBES (2010) Intergovernmental science-policy platform
for biodiversity and ecosystem services. http://IPBES.net Keywords
Larigauderie A, Mooney HA (2010) The intergovernmen-
tal science-policy platform on biodiversity and ecosys-
tem services: moving a step closer to an IPCC-like Active transport; ATP; ATP synthesis; ATPase;
mechanism for biodiversity. Curr Opin Environ Sus- Chemical energy; Electrical potential; Enthalpy;
tain 2:1–2 Entropy; Energy; Osmotic work; Photosynthesis;
Millennium Ecosystem Assessment (2005) http://www.
Proton motive force; Radiation; Respiration
maweb.org
Naeem S, Bunker D, Hector A, Loreau M, Perrings
C (2009) Biodiversity, ecosystem functioning and eco-
system services. Oxford University Press, Oxford Definition
Scholes RJ et al (2008) Toward a global biodiversity
observation system. Science 321:1044–1045
Wilson EO (ed) (1988) Biodiversity. National Academy Bioenergetics is the part of biochemistry dealing
Press, Washington, DC with ▶ energy flow through living systems. Life
Bioenergetics 271
is dependent on energy transformation reactions. operative in the biosphere. While organisms give
The ability to harness energy from a variety of just as much energy as they gain, the entropy
metabolic pathways is a property of all living associated with the energy released must exceed
organisms. the entropy associated with the energy taken
up. Organisms can develop if they interpose B
themselves into a gradient of entropy between
Overview incoming and outgoing energies.
Entropy is absorbed by live systems in the
Life and Energy form of chemical substrates and radiation. In a
Life implies work. All life systems perform work: chemical process, in ideal conditions, the sum of
chemical work is needed for the synthesis of the entropies of the reacting chemical system and
macromolecules, osmotic work is required for the environment remains constant all the time. In
the maintenance of cellular concentrations, and ideal conditions, the process is carried out revers-
electrical work is necessary for the generation of ibly, but a small change in the parameters, for
▶ proton motive force. Energy is the capacity to example, concentration, could reverse the direc-
do work. Until the principle of the conservation tion of the process.
of energy, the first law of thermodynamics, was In living systems, work generated from chem-
enunciated in the middle of the nineteenth century, ical substrates is not produced by way of heat.
progress in bioenergetics was extremely slow. Chemical energy is transformed into other forms
The second law of thermodynamics estab- of energy (e.g., ▶ Proton Motive Force, ▶ ATP)
lishes that in an isolated system, entropy without passing through a heat stage. The small
(a measure of molecular disorder) can never gradients of temperature that might exist within
decrease. The second law was enunciated for organisms are never used for work. In general,
inanimate matter; however, a violation of this live systems work isothermally and isobarically.
principle by living organisms has never been In bioenergetics, the concept of free energy is
observed. Therefore, on empirical grounds, the very useful. The molar free energy, G, is defined
validity of the second and first laws of thermody- as
namics for living organisms is generally
accepted. Nevertheless, it must be kept in mind G ¼ H TS
that living beings are open systems.
Not all processes permitted by the second law where H is the molar energy content, T is the
of thermodynamics can be used by living sys- temperature, and S the molar entropy content.
tems, for example, they cannot use the differ- Strictly, in an isobaric system, H should be called
ences in temperature generated by combustion. enthalpy and G free enthalpy. But in isobaric
The source of energy to perform biological work conditions, the volume changes are so small that
is chemical, and only few classes of chemical the difference between enthalpy and energy dis-
reactions can be used for these purposes. appears. The energetics of the reaction can be
The Earth is in a steady state. The intake of described in terms of the differences between
energy is just compensated by its loss. Energy products and reactants as
intake and energy loss occur entirely through
radiation. In a steady state, the entropy content DG ¼ DH TDS
of the Earth must be also constant. Entropy is
gained in the form of solar radiation and loss as The free enthalpy change DG has a negative
electromagnetic radiation. Entropy is also pro- value when the reaction is written in the direction
duced through irreversible processes. The rate in which it proceeds spontaneously, i.e., an exer-
of production of entropy must be equal to the gonic reaction. In the opposite direction, the reac-
net loss. Thus, in balance, the Earth emits tion is endergonic, and not spontaneous. The
entropy, and this entropic compensation is also maximum work that can be obtained from a
272 Bioenergetics
chemical reaction is given by DG, and it is reactions cannot proceed fully reversible if they
obtained when the reaction is performed revers- have finite velocities. Metabolism cannot be too
ibly. But when a reaction has an irreversible slow. Therefore, full reversibility cannot be
component, some of the energy is dissipated as attained and entropy must be generated. Hence,
heat and then additional entropy is generated. entropy is certainly generated when organisms do
While DH is mostly measured by calorimetry, external work.
DG and TDS are normally derived from measure- Experimental work shows that antagonistic
ments of chemical equilibrium. DG is related catabolic and anabolic processes always proceed
with the equilibrium concentrations of products along different pathways. Catabolic reactions are
(cp1, cp2, . . .) and reactants (cr1, cr2, . . .) and their usually exergonic; thus, the corresponding ana-
actual concentrations (c p1, c p2. . .., c r1, c r2. . ..): bolic reactions cannot proceed spontaneously, at
least not in the same conditions. Therefore, dif-
ferent pathways, which proceed at the expense of
cp1 cp2 . . . c p1 c p2 . . .
DG ¼ RTln þ RTln additional energy, are used for anabolism. In
cr1 cr2 . . . cr1 cr2 . . . general, more energy in useful form is needed
for anabolism than the energy obtained in catab-
provided that activities can be replaced by con- olism. Often, no useful energy is obtained at all in
centrations. For physicochemical standard the catabolic reactions of dynamic states. In this
conditions (pH value 0), the value DG is called case, energy is dissipated as heat.
DG . For physiological standard conditions
(pH value 7), DG is by definition DG0 . Active Transport
The equilibrium constant K is defined as One of the most important dynamic states of the
cell is the so-called active transport. Active trans-
cp1 cp2 . . . port is related with the transport of molecules and
K¼
cr1 cr2 . . . ions across the ▶ cell membrane against a con-
centration gradient. It requires osmotic work. To
Therefore, for standard conditions, perform osmotic work, free energy is required.
When the concentration of a solute in one side of

DG ¼ RTln K the membrane is different from the concentration
in the other side, diffusion to equilibrate the con-
and centrations in both sides of the membrane occurs.
The direction of the transport is given by the

DG 0 ¼ RTln K 0 change in free energy:
As DG (or DG 0 ) is a measure of the tendency a2

DG ¼ RTln
for a reaction to occur in standard conditions, it is a1
also known as the reaction affinity. The greater
the affinity, the more exergonic the reaction and a being the thermodynamic activity, which as we
the more negative DG (or DG 0 ). It has to be mentioned before in most cases can be approxi-
underlined that the exergonicity or endergonicity mated to concentration. In diffusion DG is nega-
of a reaction, in a given condition, is not deter- tive. In active transport DG is positive; this is the
mined by the standard values (DG or DG 0 ) but reason why osmotic work must operate to allow
the values of DG. The work obtainable from a the solute to cross the membrane against a con-
reaction is influenced by the actual concentra- centration gradient. If the solutes have a charge
tions, more exactly activities, of the reactants (ions), then an additional term related with the
and the products. electrical potential must be considered.
As mentioned in the ideal case of reversible Through active transport, large differences in
reactions, no entropy is generated. However, concentration of molecules (i.e., glucose) and
Bioenergetics 273
ions (i.e., H+, Na+) can be set up between the sides of the membrane. The difference in poten-
cytoplasm and the environment or between dif- tial energy (chemical energy) between the elec-
ferent cell compartments in eukaryotic cells. tron donor and the acceptor is transformed in
Obviously if a difference in concentration across proton motive force, a universal storage system
the membrane is generated by active transport, of cellular energy. B
diffusion will operate in the opposite direction; In ▶ photosynthesis, the photosynthetic reac-
thus, work must be performed all the time to tion center is excited (oxidized) by radiation, and
maintain a steady state. Solutes are pumped the excited electron is trapped by a constituent of
across the membrane by specific mechanisms. an associated electron transport chain, and as in
Specificity is extremely important in active trans- the respiration, the passage of the excited electron
port to ensure that the cell invests energy in through the different components of the chain can
transporting useful compounds. As has been promote (i) the translocation of protons produc-
pointed out, dynamic states are very important ing a gradient of protons (proton motive force) or
for life systems. No organism is known to be able (ii) the generation of cellular reducing power,
to develop without dynamic states. The complex depending of the type of photosynthesis. In the
mechanisms to maintain and regulate dynamic first case, the same excited electron goes back to
states are rather universal. One of the biggest the oxidized reaction center to reduce it to the
surprises that is emerging from the analysis of ground state (cyclic ▶ anoxygenic photosynthe-
whole-genome sequences is the high percentage sis). In the second case, an electron donor must
of genomic information devoted to active trans- donate an electron to the reaction center to bring
port, which underlines its importance. it to the ground state (oxygenic photosynthesis,
Active transport is also the basis of cell bioen- noncyclic anoxygenic photosynthesis), getting in
ergetics. Most ▶ energy conservation reactions both cases ready for another excitation reaction
are directly or indirectly related with active trans- using radiative energy. In the photosynthesis, the
port systems. From an energetic point of view, source of energy is radiation and is transformed
active transport can be classified as transport through a more or less complex system
reactions that generate useful energy for the cell (depending on the type of photosynthesis) in cel-
(primary active transport) and those that require lular energy (proton motive force and/or reducing
energy for its functioning (secondary active power).
transport). Examples of primary active transport Fermentation is the only bioenergetic system
are ▶ respiration, ▶ photosynthesis, the transport that does not make use of an electron transport
of the fermentation products, and the ▶ ATPase chain to conserve energy (see ▶ Fermentation);
activity, all of them coupled to the generation of instead cytoplasmic soluble enzymatic reactions
▶ proton motive force. Examples of secondary are able to generate cellular useful energy (ATP)
active transport are the transport of nutrients to from chemical energy (reduced carbon sub-
the cell, the maintenance of homeostatic ionic strates). Interestingly enough, because fermenta-
concentrations inside and outside of the cell, the tion is the main source of energy for these
bacterial flagellar movement produced, and the organisms, an important amount of substrate
synthesis of ATP, all of them coupled to the must be fermented for growth, and as a conse-
dissipation of the ▶ proton motive force. quence, a high concentration of fermentation
In respiration (see ▶ Respiration), an electron products are produced (i.e., ethanol, organic
donor gives electrons to an appropriated electron acids). If the organism is able to couple the trans-
acceptor through the use of an electron transport location of the fermentation products to the gen-
chain located in the ▶ cell membrane in prokary- eration of a proton gradient, then the potential
otes and in the mitochondrial membrane in energy stored as a concentration gradient of fer-
eukaryotes. The passage of electrons through mentation products, which is not useful to fuel
the chain promotes the translocation of protons, energy requiring cellular reactions, is
generating a gradient of protons between both transformed in proton motive force, a useful
274 Bioenergetics
cellular energy source. In the absence of coupling active transport required to pump sodium out of
between both transports, only diffusion applies the cell can be performed using the energy stored
and the potential energy is lost. It has been cal- as proton motive force through the coupling of
culated that up to one third of the cellular energy both transport systems.
used by fermentation organisms can be generated But energy stored in the form of proton motive
by this peculiar active transport system. force can be used for other cellular functions
All these are examples of primary active trans- different than transport, like the bacterial move-
port systems because they can transform different ment of flagella. In this case the dissipation of the
▶ energy sources in proton motive force. Once proton motive force is used to perform a mechan-
there is enough energy stored as proton motive ical work: the rotation of the flagella or the
force, it can be used to promote reactions that reverse transport of electrons in the electron
require work. The best-known example is the transport chain to generate reducing power for
translocation of cellular substrates (i.e., glucose) microorganisms that need it and cannot produce
to the interior of the cell against a concentration it by other means (many chemolithoautotrophs).
gradient. Normally, organisms, especially pro- A special case of active transport that requires
karyotes, live in habitats with extremely low con- attention is related with a membrane enzymatic
centration of useful substrates to obtain energy. complex involved in the synthesis and hydrolysis
The required concentrations of these substrates of ATP (see ▶ ATPase and ▶ ATP Synthase).
inside the cell for metabolic reactions to proceed The membrane-bound synthesis of ATP from
are several orders of magnitude higher than their ADP and phosphate is an endergonic reaction
concentration in the environment. Thus, very that requires energy. This reaction, named
active transport system is required to concentrate ▶ ATP synthase, can be fuelled by the use of
these substrates inside the cell. To do this osmotic the energy stored as proton motive force. In this
work, energy in the form of proton motive force is case, it will perform as a secondary active trans-
used. In this case, the translocation of a specific port system. But the same enzymatic complex
substrate is coupled to the dissipation (transport) can perform in the opposite direction: hydrolyz-
of proton motive force. ing ATP and using the released energy to promote
Similarly, proton motive force can be used to the translocation of protons outside the mem-
maintain a high concentration of K+ or a low brane, generating proton motive force. In this
concentration of Na+ inside the cell. Due to sim- case it is performing as a primary active transport
ple diffusion, the high concentration of potassium system. The reversibility of this reaction is the
stored inside the cell and required for an optimal core of the bioenergetic regulation of prokaryotic
performance of the different functional enzy- cells. Experimentally, it can be shown that there
matic activities licks out because the concentra- is a linear relationship between the intracellular
tion of this cation is rather low outside of the concentration of ATP and the proton motive
cells. To maintain a high concentration of potas- force. If the proton motive force measured as
sium in the cytoplasm, an active transport is membrane potential corresponds to the concen-
required, because it has to be performed against tration of ATP measured inside the cell, no activ-
a concentration gradient. The coupling between ity can be detected. If the proton motive force is
the translocation of potassium to the interior of lower than the value that will correspond to a
the cell with the dissipation of the proton motive given concentration of ATP, then ATP is hydro-
force allows maintaining the optimal cellular lyzed to generate proton motive force until both
potassium concentration. A similar situation can systems are equilibrated. If the proton motive
be found with the sodium concentration. In this force is higher than the value that should corre-
case, the optimal cellular condition is a lower spond to the ATP concentration, then the proton
concentration of this cation in the cytoplasm motive force is dissipated to increase the ATP
than in the environment. In this case, simple concentration until both systems reach equilib-
diffusion licks sodium to the cytoplasm, so the rium. With only one metabolic activity,
Biofilm 275
prokaryotes can regulate in an efficient manner with other stars and the atmosphere on extrasolar
and with little genomic investment the bioener- worlds. Astrobiology 7:252–274
Krah A, Pogoryelov D, Meier T, Faraldo-Gómez JD
getics of the cell. (2010) On the structure of the proton-binding site in
the Fo rotor of chloroplast ATP Synthases. J Mol Biol
395:20–27 B
See Also K€uhlbrandt W (2004) Biology, structure and mechanism
of P-type ATPases. Nat Rev Mol Cell Biol 5:282–295
Madigan MT, Martinko JM, Dunlap PV, Clark DP
▶ Anoxygenic Photosynthesis (2008) Brock biology of microorganisms, 12th edn.
▶ ATP Benjamin Cumming, San Francisco
▶ ATPase Mulkidjanian AY, Makarova KS, Galperin MY, Koonin
EV (2007) Inventing the dynamo machine: the evolu-
▶ ATP Synthase tion of the F-type and V-type ATPases. Nat Rev
▶ Cell Membrane Microbiol 5:892–899
▶ Electrochemical Potential Mulkidjanian AY, Galperin MY, Makarova KS, Wolf YI,
▶ Energy Koonin EV (2008) Evolutionary primacy of sodium
bioenergetics. Biol Direct 3:13
▶ Energy Conservation Nelson DL, Cox MM (2009) Lehninger principles of bio-
▶ Energy Sources chemistry, 5th edn. WH Freeman, New York
▶ Metabolism Nicholls DG, Ferguson SJ (2002) Bioenergetics 3. Aca-
▶ Mitochondrion demic, London
Noji H, Yasuda R, Yoshida M, Kinosita K Jr (1997) Direct
▶ Oxidation observation of the rotation of F1-ATPase. Nature
▶ Photosynthesis 386:299–302
▶ Proton Motive Force Pennazio S (2008) Photosynthesis: the years of light. Riv
▶ Proton Pump Biol 101:443–462
Renger G, Kuhn P (2007) Reaction pattern and mecha-
▶ Reduction nism of light induced oxidative water splitting in pho-
▶ Respiration tosynthesis. Biochim Biophys Acta 1767:458–471
Stephan E, Giovanni F, Francis-André W (2008) The
dynamics of photosynthesis. Annu Rev Genet
42:463–515
References and Further Reading Voet D, Voet JG (2004) Biochemistry, 3rd edn. Wiley,
New York
Allen JF, Martin W (2007) Evolutionary biology: out of White D (1999) The physiology and biochemistry of pro-
thin air. Nature 445:610–612 karyotes, 2nd edn. Oxford University Press, New York
Bryant DA, Frigaard NU (2006) Prokaryotic photosynthe- Xion J, Bauer CE (2002) Complex evolution of photosyn-
sis and phototrophy illuminated. Trends Microbiol thesis. Annu Rev Plant Biol 53:503–521
14:488–496 Xiong J (2006) Photosynthesis: what color was its origin?
Buch-Pedersen MJ, Pedersen BP, Veierskov B, Nissen P, Genome Biol 206:1465–6914
Palmgren MG (2009) Protons and how they are
transported by proton pumps. Eur J Physiol
457:573–579
Gr€uber G, Marshansky V (2008) New insights into
structure-function relationships between archeal ATP Biofilm
synthase (A1A0) and vacuolar type ATPase (V1V0).
Bioessays 30:1096–1109
Heathcote P, Fyfe PK, Jones MR (2002) Reaction centers: Jana Kvı́derová
the structure and evolution of solar power. Trends Institute of Botany, Academy of Sciences of the
Biochem Sci 27:79–87 Czech Republic, Trebon, Czech Republic
Itoh H, Takahashi A, Adachi K, Noji H, Yasuda R,
Yoshida M, Kinosita K Jr (2004) Mechanically driven
ATP synthesis by F1-ATPase. Nature 427:465–468
Kiang NY, Siefert J, Govindjee, Blankenship RE (2007a) Keywords
Spectral signatures of photosynthesis. I. Review of
Earth organisms. Astrobiology 7:222–251
Adaptation; Biofilm; Biomarkers; Community;
Kiang NY, Segura A, Tinetti G, Govindjee, Blankenship
RE, Cohen M, Siefert J, Crisp D, Meadows VS (2007b) Extreme environment; Microorganisms; Species
Spectral signatures of photosynthesis. II. Coevolution composition; Structure
276 Biofilm
Synonyms
Microbial mats; Periphyton
Definition
Biofilm is a layer of microorganism(s) or micro-

bial communities growing on a solid surface,
usually of thicknesses ranging between several
mm to several mm. The thicker biofilms are also
referred as ▶ microbial mats.
Overview
Biofilm communities are found in various aquatic

and subaerial ecosystems or even in the human
body (e.g., dental plaque) at liquid-solid or aerial-
solid interfaces (Fig. 1). Typical biofilms develop
on stones in streams and rivers or at the bottom of
lakes. They participate in nutrient and energy
cycling in the given ▶ ecosystem and could Biofilm, Fig. 1 Biofilms in the acidic waters of Rı́o
Tinto, SW Spain
serve as a substrate for colonization by other
species. Repeated sedimentation of inorganic
material on the growing biofilm leads to layered
structure resembling stromatolites (Krumbein possible that microorganisms within biofilms
et al. 2003). communicate by chemical signals. Such com-
Biofilms are usually formed by various munication has been detected in biofilms of
microorganisms that include autotrophs as well pathogenic bacteria, but in natural communities
as heterotrophs, eukaryotes, bacteria, and, in the very little is known of this phenomena (Kellner
most extreme conditions, archaea (Aguilera and Surette 2006).
et al. 2007a). The organization level of the bio- The development of a biofilm starts by accu-
film species can range from unicellular flagel- mulation of amorphous particles containing inor-
lates to multicellular filaments (Fig. 2). If the ganic grains and bacteria. The cells divide,
biofilm is formed by several layers of microor- produce extracellular polymeric substances,
ganisms, gradients of physical and chemical form microcolonies, and provide substrate for
factors (e.g., light, pH, O2) are established and further colonization by fungi and small eukary-
influence its structure and species composition. otic heterotrophs like amoebas. The flagellates
The microenvironment within the mature bio- also participate at the initial formation. The ses-
film could be different from that of the surround- sile genera are observed later and could require
ings and could provide protection to more establishment of some organic matrix to be
sensitive species (Ferris et al. 2005). For exam- attached on. Filamentous microorganisms are
ple, acidophilic red alga Cyanidium sp. is more last to appear. Small pieces of mature biofilm
tolerant to pH values above 3 than to those detach either due to internal signals or external
below 1, probably due to adaptation to elevated factors, like water flow, and could colonize a new
pH in biofilm interior (Kvı́derová 2012). It is surface (Aguilera et al. 2007b).
Biofilms 277
▶ Microorganism
▶ Rio Tinto
▶ Stromatolites
B
Aguilera A, Souza-Egipsy V, Gómez F, Amils R (2007a)

Development and structure of eukaryotic biofilms in
an extreme acidic environment, Rı́o Tinto (SW,
Spain). Microb Ecol 53:294–305
Aguilera A, Amaral-Zettler L, Souza-Egipsy V, Zettler E,
Amils R (2007b) Eukaryotic community structure
from Rı́o Tinto (SW, Spain), a highly acidic river. In:
Seckbach J (ed) Algae and cyanobacteria in extreme
Biofilm, Fig. 2 Example of a photosynthetic biofilm environments. Springer, Dordrecht
consisting in the association of unicellular red alga Ferris MJ, Sheehan KB, K€ uhl M, Cooksey K,
Cyanidium caldarium, filamentous green alga Wiggleswoth-Cooksey B, Harvey R, Henson JM
Klebsormidium sp., and pennate diatom Pinnularia (2005) Algal species and light microenvironment in a
sp. from Rı́o Tinto, Spain low-pH geothermal microbial mat community. Appl
Environ Microbiol 71:7164–7171
Gómez F, Walter N, Amils R, Rull F, Klingelhöfer AK,
Biofilms are the new microbial ecology frontier. Kvı́derová J, Sarrazin P, Foing B, Behar A, Fleischer I,
Biofilms have not been studied in a systematic way Parro V, Garcia-Villadangos M, Blake D, Martin
until recently due to lack of appropriate methodol- Ramos JD, Direito S, Mahapatra P, Stam C,
Venkateswaran K, Voytek M (2011) Multidisciplinary
ogies. Confocal microscopy and in situ hybridiza-
integrated field campaign to an acidic Martian Earth
tion techniques allow to study intact biofilms and analogue with astrobiological interest: Rio Tinto. Int
appreciate their complex structural diversity. For J Astrobiol 10:291–305
evaluation of photochemical performance of auto- Kellner L, Surette MG (2006) Communication in bacteria:
an ecological and evolutionary perspective. Nat Rev
trophic biofilms, methods using variable chloro-
Microbiol 4:249–258
phyll fluorescence are applied in field (Gómez Kral T, Bekkum C, McKay C (2004) Growth of
et al. 2011; Marteinsson et al. 2013) and in detailed methanogens on a Mars soil simulant. Orig Life
laboratory (Kvı́derová 2012) studies. Evol Biosph 34:615–626
Krumbein WE, Paterson DM, Zavarzin GA (2003) Fossil
An important part of microbial life on Earth is
and recent biofilms: a natural history of life on earth.
associated to biofilms. Biofilms and microorgan- Kluwer Academic Publishers, Dordrecht
isms from ▶ extreme environments are consid- Kvı́derová J (2012) Photochemical performance of the
ered interesting astrobiological model systems acidophilic red alga Cyanidium sp. in a pH gradient.
Orig Life Evol Biosph 42:223–234
(Aguilera et al. 2007b; Gómez et al. 2011; Kral
Marteinsson V, Vaishampayan P, Kvı́derová J, Mapelli F,
et al. 2004), specially as biomarkers of extinct or Medori M, Calfapietra C, Aguilera A, Hamisch D,
extant life in habitable planets like Mars or on Reynisson E, Magnússon S, Marasco R, Borin S,
Galilean moons. Calzada A, Souza-Egipsy V, González-Toril E,
Amils R, Elster J, Hänsch R (2013) A laboratory of
extremophiles: Iceland Coordination Action for
Research Activities on Life in Extreme Environments
See Also (CAREX) field campaign. Life 3:211–233
▶ Colonization, Biological
▶ Concentration Gradients
▶ Ecosystem Biofilms
▶ Microbial Mats ▶ Microbial Mats
278 Biogenicity
symmetry, and isotopic composition of chemical

Biogenicity traces of life; and (3) the environmental distribu-
tion or geological context of traces of life. These
Nicola McLoughlin three approaches to investigating the biogenicity
Department of Earth Science and Centre for of candidate traces of life are mutually
Geobiology, University of Bergen, Bergen, reinforcing and should be accompanied by efforts
Norway to falsify potential abiotic explanations for the
observations. The so-called ▶ dubiofossils or
▶ pseudofossils are features that are highly ques-
Keywords tionable traces of life and more likely explained
by abiotic processes that can mimic life
Biosignatures and traces of life; History and ori- (Hofmann 1971).
gins of life; Life detection Specific biogenicity criteria have been tailored
for the different classes of biosignatures found on
Earth. For ▶ microfossils in the rock record,
Synonyms biogenicity criteria have been proposed by
Buick (1990) and Brasier et al. (2004), and in
Biosignature summary these focus on the size distribution of
the population; morphological features such as
branching, septation, evidence for cell walls,
Definition nuclei, or extracellular polymeric substances;
their orientation, especially evidence of colonial
Biogenicity refers to any chemical and/or mor- behavior, tiering, or phototaxis; and lastly evi-
phological signature preserved over a range of dence of primary environmentally controlled dis-
spatial scales in rocks, minerals, ice, or dust par- tribution and/or subsequent decay. The
ticles that are uniquely produced by past or pre- importance of such criteria has been highlighted
sent organisms. This includes elemental and by abiotic experiments that produce microfossil-
isotopic signatures diagnostic of life, which can- like filamentous biomorphs in the laboratory
not be formed by purely abiotic processes. These (Garcia-Ruiz et al. 2003). These microfossil
may be accompanied by textural remains with biogenicity criteria have also been adapted for
shapes, orientations, and abundances that meteorite samples, especially for those purported
uniquely result from the growth or decay of to contain fossilized ▶ magnetobacteria
(once) living organisms. Further support for (Thomas-Kerpta et al. 2001).
biogenicity can be shown if the distribution and For laminated sedimentary structures known as
abundance of this evidence is controlled by bio- ▶ stromatolites that are formed by the interaction
logically significant primary variables such as of ▶ microbial mats and sediments, biogenicity
light, temperature, and nutrient gradients. criteria have been advanced by Buick
et al. (1981) and contrasted with abiotic artifacts
by McLoughlin et al. (2008). A primary sedimen-
Overview tary origin along with complex laminated macro-
morphologies such as domes, columns, and
Biogenicity criteria are used to assess the likeli- branches is supportive of a biological origin, but
hood of a biological origin for candidate traces of it is the micro-fabrics of stromatolites, which,
life. Biogenicity criteria address observable and preservation permitting, can be one of the best
quantifiable features that can be broadly divided indicators of a biological origin (Hofmann 2000).
into three types: (1) the morphological complex- For the related phenomenon of wrinkle mat
ity and size distribution of textural traces of life; textures, also termed ▶ microbially induced sedi-
(2) the elemental composition, bonding, mentary structures, biogenicity criteria have been
Biogeochemical Cycles 279
formulated by Noffke (2009). Lastly, for micro- microborings on early Earth and beyond. Astrobiology
cavities produced by rock-dwelling organisms that 7:10–26
McLoughlin N, Wilson LA, Brasier MD (2008) Growth of
tunnel into rock substrates including carbonates synthetic stromatolites and wrinkle structures in the
and volcanic glass, biogenicity criteria have been absence of microbes – implications for the early fossil
developed by McLoughlin et al. (2007). record. Geobiology 6:95–105 B
Noffke N (2009) The criteria for the biogenicity of
microbially induced sedimentary structures (MISS)
in Archean and younger, sandy deposits. Earth Sci
See Also Rev 96:173–180
Thomas-Kerpta KL, Clemett SJ, Bazylinski DA,
▶ Archean Traces of Life Kirschvink JL, McKay DS, Wentworth SJ, Vali H,
Gibson JEK, McKay MF, Romanek CS (2001) Trun-
▶ Biomarkers cated hexa-octahedral magnetite crystals in
▶ Biomarker, Isotopic ALH84001: presumptive biosignatures. Proc Natl
▶ Biomarkers, Morphological Acad Sci U S A 98:2164–2169
▶ Dubiofossil
▶ Endogenicity
▶ Endolithic
▶ Fossil Biogeochemical Cycles
▶ Life
▶ Magnetotactic Bacteria David C. Fernández-Remolar
▶ Microbial Mats Centro de Astrobiologı́a (INTA-CSIC), INTA,
▶ Microfossils Torrejón de Ardoz, Spain
▶ Pseudofossil
▶ Stromatolites
▶ Syngenicity Keywords
Biogeochemistry; Biosphere; Earth systems;

References and Further Reading Ecosphere; Geobiology; Habitability; Homeosta-
sis; Hydrosphere; Matter; Energy fluxes
Brasier MD, Green OR, McLoughlin N (2004) Character-
ization and critical testing of potential microfossils
from the early Earth: the Apex ‘microfossil debate’
and its lessons for Mars sample return. Int J Astrobiol Synonyms
3:139–150
Buick R (1990) Microfossil recognition in Archaean Nutrient cycles
rocks: an appraisal of spheroids and filaments from
3500 M.Y old chert-barite at North Pole, Western
Australia. Palaios 5:441–459
Buick R, Dunlop JSR, Groves DI (1981) Stromatolite Definition
recognition in ancient rocks: an appraisal of irregularly
laminated structures in an early Archean chert-barite
unit from North Pole, Western Australia. Alcheringa
The biogeochemical cycles is a theoretical concept
5:161–181 that describes the transfer of matter and energy
Garcia-Ruiz JM, Carnerup AM, Christy AG, Welham NJ between the ▶ biosphere and the other active res-
(2003) Morphology and ambiguous indicator for ervoirs of Earth like the atmosphere, ▶ hydro-
biogenicity. Astrobiology 2:353–369
sphere, and lithosphere (Schlessinger 1997). The
Hofmann HJ (1971) Precambrian fossils, pseudofossils,
and problematica in Canada. Bull Geol Surv Can main processes can be understood as the storage or
189:146 release of energy by the biosphere through the
Hofmann HJ (2000) Archean Stromatolites as microbial uptake or liberation of those molecular species
Archives. In: Riding RE, Awramik SM (eds) Microbial
bearing essential components, the so-called nutri-
sediments. Springer, Berlin
McLoughlin N, Brasier MD, Wacey D, Green OR, Perry ents, for life. The main biogeochemical cycles
RS (2007) On biogenicity criteria for endolithic involve different molecular species whose
280 Biogeochemical Cycles
composition provides the essential elements for numerous nutrients, it benefits from the many
both building polymeric structures and the bio- fluxes of matter and energy sourced in some
chemical transduction of energy in cells, like C, Earth systems, which transport energy and matter
N, S, P, and O. The biogeochemical cycles are to the biosphere. Such planetary processes would
activated when the biosphere couples to the nutri- not have been adopted over long-term periods,
ent fluxes that are supplied from specific sites or which in some cases show homeostatic proper-
active centers on Earth (e.g., volcanic centers, ties, without long-lasting nutrient recycling over
hydrothermal systems) that are essentially energy geological time. As a consequence, the biogeo-
sources which maintain the supply of matter and chemical cycles make up the global network that
energy to the Earth surface. The biosphere, as the interconnects the biotic and abiotic systems to
top-level ecosystem, has its own energy trans- other essential elements of Earth biogeochemis-
ducers, a role played by the primary producers try. Such components with their different func-
that occupy the different habitats of the planet. tions, structures, and distributions on Earth are
They promote the transport of energy in the form indeed cornerstones for the cyclical emergence
of molecular compounds from the lithosphere to and maintenance by inflow and outflow of matter
the different reservoirs of the Earth (Lovelock and energy in the form of nutrient exchange. The
2000). Most of these active systems are powered essential unit in biogeochemistry is the reservoir
by the release of the inner heat of Earth, which, in that corresponds to the storing component for
the form of aerial or subaqueous systems, intro- molecular nutrient-bearing species. In this
duces at the regional or global scale different vol- sense, the four main reservoirs on Earth
atiles and other chemical species usable by living (atmosphere, hydrosphere, biosphere, and litho-
forms (Schlessinger 1997). Matter and energy sphere) are capable of harvesting nutrients from
cycling is accomplished when the Earth fluxes elements with different redox and chemical state.
drive the nutrients and other components to spe- The framework of reservoirs is strongly
cific sinks where they are generally recycled by constrained by the inflow and outflow of the nutri-
biochemical or geochemical-mediated reactions to ent through critical locations where the energy is
compounds having a different chemical state. One collected, transformed, or derived to another res-
typical example of this process is the biogeochem- ervoir. Some of these critical sites, or active cen-
ical cycling of sulfur in its highest oxidized state, ters, have been considered to serve as planetary
SO42 to more reduced compounds, which are organs (Lovelock 2000), whose disintegration
sourced in volcanic centers or precipitated would collapse the energy flows that supply the
throughout the hydrothermal systems as metallic biosphere. In this sense, the biosphere has pro-
sulfides (e.g., Fe2S), hydrogen sulfide (H2S), or vided itself with some of them in the form of
dioxide of sulfur (SO2). All these molecular spe- specific trophic groups led by autotrophic organ-
cies can be potentially used by organisms as elec- isms that occupy specific habitats. In the bio-
tron donors to obtain energy, as well as raw sphere, the energy collectors supply nutrients to
material to produce sulfur-bearing machinery the rest of their biological components, which are
under microbial assimilation (Fenchel et al. 1998). stored in the biosphere in the form of biomole-
cules. However, the nutrient and energy supply is
not only located in the biogeochemical active cen-
Overview ters of the biosphere but also in some biologically
independent systems that are essentially associ-
The biogeochemical cycles can be considered a ated to heat dissipation centers. A good example
complex network of interrelated mechanisms by of these are the volcanic centers that are responsi-
which the biosphere globally interacts with the ble for introducing huge quantities of key nutrients
different reservoirs through very complex, posi- for the biosphere in the form of volatiles (e.g.,
tive and negative, feedback-controlled reactions H2S, SO2, CO2, CH4, H2, NH3, NO2, etc.) into
(Margaleff 1968). As the biosphere requires the atmosphere or, after their transportation, into
the hydrosphere (Fig. 1). They are eventually inte- Interestingly, the introduction of sulfur-bearing
grated by the biosphere through direct uptake of compounds into the atmosphere and their subse-
gases or after their dissolution into a fluid phase to quent oxidation into sulfate has been found to be a
be used as raw material for building biomolecules key mechanism in nucleating clouds, which has a
or as an energy source. planetary impact as it increases the albedo and, B
The activity of the biosphere can precipitate, therefore, decreases the solar radiation affecting
biomineralize, or release a wide range of inor- the Earth’s surface (Charlson et al. 1987). As a
ganic compounds bearing nutrients such as sulfur, consequence, sulfur cycling by phytoplankton is
carbon, or phosphorous (Sundquist and Visser an essential feedback mechanism to control to
2003; Ruttenberg 2005) that can potentially pro- some extent climatic conditions in the planet.
vide chemical energy to the biosphere itself (e.g., There are other nutrients that are essential for
sulfide, ammonia, or hydrocarbon bio-oxidation) life in very low concentrations and are released
or raw material to the other reservoirs (Fig. 1). back to the atmosphere following the pathways
They can be exchanged as simple volatile similar to that of dimethyl sulfide. This is the case
exchange, anionic flow or biosedimentation of iodine that is released into the atmosphere by
toward the other three reservoirs that, after their marine phytoplankton in the form of different
chemical transformations, can then be recycled by volatile compounds such as methyl iodide
the biosphere. Such processes result in the elimi- (Butler et al. 1981; Manley and de la Cuesta
nation of nutrients from the biosphere through 1997), which is decomposed by photolysis to
specific sites or sinks where matter and energy more oxidized forms of iodine under reaction
are driven to the other Earth reservoirs. One par- with oxidants in the troposphere. The iodine is
adigmatic example of nutrient loss from the bio- returned back to the lithosphere or hydrosphere
sphere into the lithosphere is limestone by rainfall or simple influx from the atmosphere
production by unicellular planktonic algae and after being included in the aerosols which are
foraminifera, which biomineralize carbonate- active cloud-nucleating agents on Earth (Baker
forming shells ((Zondervan et al. 2001); Fig. 1). 2005). Interestingly, the nutrient exchange
After the microorganism’s death, the carbonatic between the different reservoirs activates some
shells continuously rain down on the ocean floor feedback mechanisms involved in maintaining
and eventually lead to the global formation of ▶ homeostasis conditions on a planetary scale
sedimentary carbonates (Fig. 1). Moreover, these (Charlson et al. 1987; Lovelock 2000).
oceanic deposits can be transported from the sed- A last party in the distribution of components is
imentary areas to the subduction zones where the the transport agents in the atmosphere, the hydro-
carbonate is transformed back into CO2 by ther- sphere, or the lithosphere, which act as natural
mal decomposition in deep areas of the litho- carriers to spread these compounds to the different
sphere (Sundquist and Visser 2003), which can reservoirs including the biosphere. Most of them
then be returned to the atmosphere through exha- are activated by simple physical gradients
lation of volatile compounds in the volcanic cen- depending on local or regional changes in temper-
ters. Some other nutrients are directly released to ature, density, pressure, or gravity. Although
the atmosphere in the form of volatilized organics, potential energy plays an essential role in fluvial
which can be released back to both hydrosphere transport, most of the transporting agents are
and biosphere once they are oxidized by reactions powered by the latitudinal difference in tempera-
driven by photochemistry. This is the case of ture that resulted from the varying incidence angle
some organic compounds like dimethyl sulfide of solar radiation on the Earth’s surface. The
(Fig. 1), one of the main sources of sulfur in the hydrological cycle on Earth provides several
biosphere, which is released into the atmosphere agents (rainfall, streams, and fluvial systems,
by phytoplankton, but is returned to the hydro- etc.) that are involved in the transportation of
sphere in the form of sulfate after being oxidized nutrients to the different Earth reservoirs involved
by UV in the atmosphere (Brimblecombe 2005). in sustaining element cycling.
282 Biogeochemical Cycles
6
UV
{−OH, H2O}
CO2 H2S SO2 H2SO4
CO2
pH = 5.2
5
7 5 CO2
5 S(CH2)3 CO2 CH2O 2 4
CO2 2 CH2O
1 CaCO3 2 CO2 CaCO3

CO2 2 CH2O CaCO3 3 O 3
CH2O 2
CO2 [CH2O] CH
SO2 CO2
CaCO3
3 CH2O 2 CO2 3
4 4 4 4 4
Carbonates
Biomass and organics
Weathered crust and sediments
1 Main carbon sources in geothermal systems through CO2 supply to the atmosphere
2 Organic carbon reservoir (biosphere and organic by-products)
3 Inorganic carbon reservoir (carbonates)
4 Inorganic and organic carbon sinks
5 Recycle of carbon back to the atmosphere from crust and ocean
6 Precursor formation for sulfuric acid (H2S oxidation through UV-mediated reactions)
7 Cloud nucleation by sulfuric acid generation
Biogeochemical Cycles, Fig. 1 Biogeochemical cycle also effective processes to remove carbon from the Earth
of carbon coupled to the sulfur cycle. Diagram shows the surface. The organic oxidation, thermal decomposition, or
main pathways and reservoirs integrating the whole car- simple carbonate dissolution through diagenesis can cycle
bon cycle and how it interplays with the sulfur cycle. back into the atmosphere (5) the carbon as carbon dioxide
Volcanic centers and other geothermal-driven systems to compensate the carbon loss through the microbial
(e.g., hydrothermal systems) release carbon to the atmo- uptake, carbonate precipitation, and/or simple weathering.
sphere (1) mainly in the form of CO2 but also CH4 and Sulfur can be oxidized to sulfuric acid through very com-
CO. The carbon is fixed as organic or inorganic carbon plex photochemical reactions starting on hydrogen sulfide
through the biosphere growth and the production of oxidation and ending on the sulfite hydration. Thus, sul-
organics (2) and the precipitation of carbonates (3), furic anion as a powerful nucleation agent of clouds
which can be produced by biomineralization, simple pre- (6) favors the supply of meteoric water to the surface
cipitation by biological mediation, oversaturation by abi- that transports C-bearing gases and other volatiles to the
otic mechanisms, and/or simple weathering. The carbon is crust, making the carbon available for its fixation as
removed from the crust by sediment burial of carbonates organic or inorganic compounds. Sulfur is released back
and organics (4); carbonate precipitations in the subsur- to the atmosphere (7) in combination with sulfur as
face by biological uptake and weathering mechanisms are dimethyl sulfide by algal activity
Basic Methodology determination of volatile fluxes in water masses,

soils, or sediments are characterized by the use of
Biogeochemistry, the discipline that studies the highly sophisticated systems that integrate the
interactions among the biosphere and the different sampling cell and the instrumentation system
Earth reservoirs, emerges from the interaction of (Heyer and Berger 2003). Gas quantification B
experimental sciences like physics, chemistry, requires pressurized chambers, which maintain
biology, and geology, which use a wide diversity the natural concentration of volatiles. On the
of methodologies and techniques. The biogeo- other hand, nutrients taking part in the composi-
chemical cycles are powered by matter and energy tion of minerals or dissolved ions can be analyzed
exchanges that are experimentally measurable by using a wide variety of techniques that can deter-
reservoir imbalance over timescales. In this sense, mine elemental compositions such as atomic
reaction kinetics plus thermodynamics, related to absorption, thermal fluorescence, mass spectrom-
energy flow and storage, are the essential back- etry, or liquid chromatography.
ground concepts in the description of how biogeo-
chemical cycles are maintained (Schlessinger
1997). Furthermore, system modeling, an essential Applications
tool deeply rooted in linear and nonlinear mathe-
matics, has not only increased our understanding The study of the biogeochemical cycles touches
the dynamics of cycling on Earth connecting the core of one of the essential questions in astro-
research on local and regional biogeochemical biology, whether a planet can sustain life through
cycling of single nutrients to other compounds the activation of the matter and energy fluxes
but also provides a context on a higher planetary between the past or modern reservoirs. The four
scale (Sellers et al. 1997). Obviously, building up main ingredients – liquid water, nutrients, energy
system models requires the collection of quantita- sources, and chemical disequilibrium – that
tive data to sustain consistent modeling. The most enable a planet to become potentially habitable
direct, but not the easiest, way to quantify fluxes are also the same key elements that initiate and
and storage of matter and energy through and in maintain the biogeochemical cycles operated by
Earth reservoirs is based on experimental mea- a hypothetical extraterrestrial biosphere. The
surements that determine the gain or loss of mea- chemical disequilibrium, among all ingredients,
surable quantities of units per year. This can be emerges as essential to fuel the matter and energy
done on local or regional scales after collecting transfer that maintains the cycling. There are
samples of air, water, sediments, soils, or rocks several attempts to visualize how biogeochemical
through fieldwork. In most of cases, specific sam- cycling could operate in modern planetary bodies
pler devices are used to recover gas, liquid, sustained by redox disequilibrium maintained
organic, or biological materials that are gathered through photochemical reactions that supply oxi-
over time in order to infer the rates of biogeochem- dizing and reducing compounds to the planetary
ical fluxes. In the case of volatiles, the analysis of surface. For Mars, a planet covered by a thin
different reservoirs interacting in a given area CO2-rich atmosphere, the radiolysis of methane
requires different in situ techniques made possible has been suggested as a mechanism to provide H2
by recent state-of-the-art analytical instruments as an electron donor for microbial life in the
such as spectrophotometers, visible and infrared subsurface (Weiss et al. 1999), the only region
spectroscopy, gas chromatography, or mass spec- in this planet that escapes the high radiation rates
trometry, among others (Viollier et al. 2003). the surface areas are exposed to. Obviously, the
However, accurate analysis for gases in geological production of reducing compounds could be the
and water samples is frequently based on careful primary source for a wider diversity in Mars
sampling processes, which require pressurized subsurface biosphere based not only on
sampling cells to collect the volatile phases under chemolithotrophic but also on heterotrophic
controlled conditions. The quantification and metabolism. Similar cycling for carbon has been
284 Biohazard Assessment Protocol
proposed for Europa (Chyba and Phillips 2001), Heyer J, Berger B (2000) Methane emission from the
an icy satellite of Jupiter with no atmospheric coastal area in the Southern Baltic Sea. Estuar Coast
Shelf Sci 51(1):13–30
shield and exposed to a heavy bombardment of Lovelock J (2000) Gaia, a new look at life on Earth,
ionized particles like Na+. However, subsurface 4th edn. Oxford University Press, Oxford
radiolysis (Blair et al. 2007) is another mecha- Manley SL, de la Cuesta JL (1997) Methyl iodine produc-
nism that has been recently been considered as a tion from marine phytoplankton production. Limnol
Oceanogr 42:142–147
provider of the chemical disequilibrium neces- Margaleff R (1968) Perspectives in ecological theory. The
sary for sustaining living forms in planetary sys- University of Chicago Press, Chicago
tems other than Earth’s. Ruttenberg KC (2005) The global phosphorous cycle. In:
Holland H, Turekian KK (eds) Treatise of geochemis-
try, vol 8. Elsevier, Amsterdam
Schlessinger WH (1997) Biogeochemistry: an analysis of
See Also global change. Academic, San Diego
Sellers PJ, Dickinson RE, Randall DA, Betts AK, Hall FG,
▶ Autotrophy Berry JA, Collatz GJ, Denning AS, Mooney HA,
Nobre CA, Sato N, Field CB, Henderson-Sellers
▶ Biosphere A (1997) Modeling the exchanges of energy, water,
▶ Carbon Cycle, Biological and carbon between continents and the atmosphere.
▶ Carbon Dioxide Science 275:502–509
▶ Gaia Hypothesis Sundquist ET, Visser K (2003) The geologic history
of the carbon cycle. In: Holland H, Turekian KK
▶ Heat Flow, Planetary (eds) Treatise of geochemistry, vol 8. Elsevier,
▶ Heterotroph Amsterdam
▶ Homeostasis Viollier E, Rabouille C, Apitz SE, Breuer E, Chaillou G,
▶ Hydrosphere Dedieu K, Furukawa Y, Grenz C, Hall P, Janssen F,
Morford JL, Poggiale J-C, Roberts S, Shimmield T,
▶ Hydrothermal Environments Taillefert M, Tengberg A, Wenzhöfer F, Witte
▶ Lithosphere, Planetary U (2003) Benthic biogeochemistry: state of the art
▶ Nitrogen Cycle, Biological technologies and guidelines for the future of in situ
▶ Nitrogen Fixation survey. J Exp Mar Biol Ecol 285–286:5–31
Weiss P, Yung YL, Nealson KN (1999) Atmospheric
▶ Sulfur Cycle energy for subsurface life on Mars? Proc Natl Acad
Sci 97:1395–1399
Zondervan I, Zeebe RE, Rost B, Riebel U (2001) Decreas-
ing marine biogenic calcification: a negative feedback
References and Further Reading on rising atmospheric pCO2. Global Biogeochem
Cycles 15:507–516
Baker AR (2005) Marine aerosol iodine chemistry: the
importance of soluble organic iodine. Environ Chem
2:295–298
Blair CC, D’Hondt S, Hondt S, Spivack AJ, Kingsley RH
(2007) Radiolytic hydrogen and microbial respiration
in subsurface sediments. Astrobiology 7:951–970 Biohazard Assessment Protocol
Brimblecombe P (2005) The global sulfur cycle. In:
Holland H, Turekian KK (eds) Treatise of geochemis-
Catharine A. Conley
try, vol 8. Elsevier, Amsterdam
Butler ECV, Smith JD, Fisher NS (1981) Influence of NASA Headquarters, Washington, DC, USA
phytoplankton on iodine speciation in seawater.
Limnol Oceanogr 26:382–386
Charlson RJ, Lovelock JE, Andreae MO, Warren SG
(1987) Oceanic phytoplankton, atmospheric sulphur,
Definition
cloud albedo and climate. Nature 326:655–661
Chyba CF, Phillips CB (2001) Possible ecosystems and Any sample brought to Earth from another plan-
the search for life on Europa. Proc Natl Acad Sci etary body is subjected to an assessment for
98:801–804
▶ planetary protection according to the recom-
Fenchel T, King GM, Blackburn TH (1998) Bacterial
biogeochemistry: the ecophysiology of mineral mendations of the ▶ COSPAR planetary protec-
cycling. Academic, San Diego tion policy. If the planetary body has the potential
Bioindicator 285
to host indigenous life, and the mission is cate- determining the carbon isotope ratio
gorized as a “restricted Earth return,” the samples (as discussed in the review of Etiope and Sher-
must be subjected to rigorous analyses and tests wood Lollar 2013). In contrast, the term
in order to prevent release of any possible extra- “biosignature” refers to a robust signal that is
terrestrial life into the Earth’s environment. strongly suggestive of life. For example, the B
A biohazard assessment protocol that has been atmospheric detection of an Earth-like amount
reviewed by an appropriate group of scientific of nitrous oxide (N2O) for an Earth-like planet
experts must be performed to determine whether orbiting a sunlike star would constitute a
extraterrestrial organisms or other hazards are biosignature because according to current knowl-
contained in materials returned from other plan- edge, such a signal could not be produced by
etary bodies. anything other than microbes. In other words,
the known atmospheric abiotic sources (for an
overview see, e.g., Kaiser and Röckmann 2005)
for this gas are much too slow to “mimic” life in
See Also
such a case.
The boundary between a bioindicator and a
▶ COSPAR
biosignature is not strictly defined. Strong
(technological) biosignatures indicative of
▶ Planetary Protection Category
advanced life have been suggested, e.g., the
detection of emitted electromagnetic waves sim-
ilar to those on Earth or the atmospheric detection
of (industrially produced) chlorofluorocarbons.
Bioindicator In such cases, it is unforeseeable as to how these
signals could arise without (intelligent) life.
John Lee Grenfell The detection of O2 (hence O3 from which it is
German Aerospace Center (DLR), Berlin, formed) would immediately initiate a debate as to
Germany the origin – biotic or otherwise – of such signals.
Any process which produces significant atomic
oxygen (O), e.g., via CO2 photolysis (potentially
Definition important in thick, Venus-like atmospheres)
and/or H2O photolysis (potentially important in
In exoplanet science the term “bioindicator” hot, steam atmospheres), can, via the self-
refers to an indication (or “hint”) that life as we reaction of O, lead to abiotic O2 production in
know it could be present – but in order to decide the atmosphere. Selsis et al. (2002) discuss these
more definitively would require additional issues. To address the question whether atmo-
information. spheric O2 and O3 could constitute a biosignature,
would therefore benefit from more theoretical
studies to improve understanding of the associ-
Overview ated range of abiotic sources.
Vegetation on Earth produces simple organic
The detection, e.g., of atmospheric methane in an molecules such as chloromethane (CH3Cl)
exoplanetary atmosphere would constitute a (Hewitt and Jackson 2009), but again each case
bioindicator because additional information is in hand would need to be investigated by apply-
needed in order to determine whether or not the ing theoretical studies adapted to the particular
source is biological. This, in the case of methane, conditions (e.g., stellar flux, spectrum, etc.) in
can be tested, e.g., by comparing the ratio of its order to determine the (in situ abiotic) gas-phase
outgassing amount with other outgassed species sources. In this process of elimination, life is the
such as H2 (see, e.g., Oze et al. 2012) and/or by “last man standing.”
286 Bioindice
See Also Definition
▶ Biosignature Bioinformatics, also called computational biol-

ogy, is a recent discipline whose aims are to
store, retrieve, classify, and organize information
about biological systems, to detect their evolu-
Etiope G, Sherwood Lollar B (2013) Abiotic methane on tionary and functional patterns, to suggest laws of
Earth. Rev Geophys 51(2):276–299 biological organization through statistical analy-
Hewitt CN, Jackson AV (eds) (2009) Atmospheric science sis, and to predict the results of new experiments
for environmental scientists. Wiley-Blackwell, by extrapolating experimental data with the help
Oxford, p 105
Kaiser J, Röckmann T (2005) Absence of isotope of evolutionary or physically inspired reasoning.
exchange in the reaction of N2O + O(1D) and the
global 17O budget of nitrous oxide. Geophys Res Lett
32:LI15808 History
Oze C, Jones C, Goldsmith JI, Rosenbauer RJ (2012) Dif-
ferentiating biotic from abiotic methane genesis in
hydrothermally active planetary surfaces. Proc Natl One can say that bioinformatics originated with
Acad Sci U S A 109:E3211 the need to compare protein sequences in an
Selsis F, Despois D, Parisot J-P (2002) Parisot, signature objective and automatic way in the early 1960s,
of life on exoplanets: can Darwin produce false posi-
tive detections? Astron Astrophys 388:985–1003 inspired by the recent discipline of information
theory, which led to the proposal of the molecular
clock in ▶ protein evolution. Since then,
sequence analysis has become more complex,
addressing the comparison of the first complete
Bioindice genomes in the 1990s. Anfinsen experiments in
the 1950s–1970s motivated the holy grail of com-
▶ Biomarkers putational biology: to predict protein structures
from physical principles (Anfinsen 1973). While
this dream did not materialize yet, it attracted
many physicists to the field and stimulated a
Bioinformatics very rich and fruitful dialogue with statistical
mechanics. Bioinformatics is nowadays a kind
Ugo Bastolla of two-headed animal, trying to combine the his-
Unidad de Bioinformática, Centro de Biologı́a torical approach of evolutionary theory with the
Molecular “Severo Ochoa,” CSIC-UAM, universality of physical laws. This hectic field is
Madrid, Spain benefiting from the open-access mentality in
which scientists all over the world freely share
their algorithms and databases through the
Keywords Internet.
Algorithms; Databases; Drug design; Gene net-

works; Macromolecules; Molecular evolution; Overview
Protein folding; Sequence analysis; Structural
bioinformatics; Systems biology The scope of bioinformatics is rapidly evolving.
Currently, its main fields are the following:
Synonyms 1. Sequence analysis: alignment and compari-

son of macromolecular sequences to predict
Computational biology structural and functional relationships;
Bioinformatics 287
algorithms for fast search of related interactions between the components of biologi-
sequences in large databases, like Blast; sta- cal systems, while mathematical modeling of
tistical models of sequence families (hidden evolutionary processes has been demonstrated
Markov models); and analysis of splice var- to have a rigorous formal analogy with statistical
iants in different tissues and organisms physics. Bioinformatics and its sister disciplines B
2. Structural bioinformatics: macromolecular are more and more central in a dialogue with
structure alignment and classification; analy- experimental biology, both as tools for predic-
sis of structural evolution; prediction of sections and information retrieval and as a concep-
ondary, tertiary (protein and RNA folding), tual framework to look for general organizational
and quaternary (protein-protein and protein- and evolutionary principles that help us to better
nucleic acid interactions) structure; predic- understand living matter.
tion of folding stability upon mutation;
prediction of the protein propensity to
misfold and form amyloid fibrils or amor-
See Also
phous aggregates, involved in diseases; anal-
ysis and prediction of conformation changes
▶ Evolution, Molecular
upon molecular binding; and prediction of
▶ Genome
naturally disordered regions of a protein
▶ Genomics
3. Computational drug design: virtual screening
▶ Nucleic Acids
of candidate drugs by predicting their affinity
▶ Phylogenetic Tree
to target ▶ proteins through protein-drug
▶ Phylogeny
docking or other techniques
▶ Protein
4. Analysis, classification, and prediction of
▶ Proteins, Primary Structure
protein function
▶ Proteins, Quaternary Structure
5. Molecular evolution: reconstruction of evo-
▶ Proteins, Secondary Structure
lutionary patterns and phylogenetic trees and
▶ Proteins, Tertiary Structure
simulations of evolutionary processes
▶ Proteome, Proteomics
6. Gene expression analysis: statistical analysis
of mRNA levels measured through
microarrays and other techniques to detect
co-regulated genes and compare different References and Further Reading
cell types and tissues in health and disease
7. Proteomic analysis to detect gene expression Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ
(1990) Basic local alignment search tool. J Mol Biol
by measuring protein content 215:403–410
8. Analysis and prediction of gene networks: Anfinsen CB (1973) Principles that govern the folding of
DNA regulation through transcription protein chains. Science 181:223–230
factors; posttranslational modifications of Bastolla U, Porto M, Roman HE, Vendruscolo M (eds)
(2007) Structural approaches to sequence evolution.
proteins, in particular phosphorylation;
Springer, Berlin/Heidelberg
protein-protein interaction networks; and Bourne PE, Weissig H (eds) (2003) Structural bioinfor-
biochemical pathways matics. Wiley-Liss, Hoboken
9. Algorithm improvement Durbin R, Eddy S, Krogh A, Mitchison G (1998) Biolog-
ical sequence analysis. Cambridge University Press,
10. Database management and integration Cambridge
11. Last but not least, literature mining to Lesk AM (2002) Introduction to bioinformatics. Oxford
retrieve information about biological sys- University Press, Oxford/New York
tems of interest Nei M, Kumar S (2000) Molecular evolution and phylo-
genetics. Oxford University Press, Oxford
Ouzounis CA, Valencia A (2003) Early bioinformatics:
Points 6–8 also belong to systems biology, an the birth of a discipline-a personal view. Bioinformat-
interdisciplinary field that focuses its study on the ics 19:2176–2190
288 Biological Efficacy
See Also
Biological Efficacy
▶ Bioburden
Catharine A. Conley ▶ Bioburden Reduction
NASA Headquarters, Washington, DC, USA ▶ Microorganism
▶ Spore
Definition
The biological efficacy of a process is used to

describe the effectiveness of a specified process
at affecting biological organisms. Efficacy refers Biological Networks
to an expected result. For instance, the biological
efficacy of the ▶ DHMR is described by the mea- Emma Hart
sured reduction of the ▶ bioburden. School of Computing, Edinburgh Napier
University, Edinburgh, UK
See Also
Keywords
▶ Bioburden
▶ DHMR
Biological network; Graph; Hub; Motif; Power
law; Scale-free network
Synonyms
Biological Evolution
Metabolic networks; Protein-protein interaction
networks; Transcription networks
Biological Indicator Definition
Catharine A. Conley A biological network is an abstract representation

NASA Headquarters, Washington, DC, USA of a biological system as a graph in which nodes
in the graph represent components in the system
(genes, cells, molecules) and links between the
Definition nodes represent interactions between compo-
nents. Links may be weighted to represent
In ▶ planetary protection, biological indicators strength of interactions. The resulting graph has
are used to establish the efficacy of a ▶ bioburden a particular topology which can be used to under-
reduction process. A well-characterized prepara- stand function.
tion of ▶ microorganisms (usually ▶ spores) is
subjected to the ▶ bioburden reduction protocol.
A number of biological indicators are commer- Overview
cially available in the form of a population of
spores generated from a species of microorgan- Recent advances in technology have resulted in
ism with known levels of resistance to a particu- an explosion in the amount of data that can be
lar bioburden reduction process. collected from biological systems. Techniques
Biological Networks 289
such as mass spectrometry, yeast two-hybrid include Saccharomyces cerevisiae (yeast),

assays, and other high-throughput methods have Helicobacter pylori (▶ bacteria), as well as
enabled significant advances in the identification those in Drosophila and humans.
of components, expression patterns, and interac- • Metabolic networks: Biochemical data has
tions within biological systems. Analyzing such enabled the construction of metabolic net- B
vast data sets is challenging. Furthermore, the works (e.g., ▶ glycolysis) which describe the
data sets are often incomplete and perhaps inac- metabolic and physical processes that deter-
curate. However, by considering the biological mine the physiological and biochemical prop-
system as a network or graph in which entities in erties of cells, usually focusing on the basic
the network are represented by nodes, with links chemical pathways that generate the essential
between nodes representing interactions, the components used in biochemical reactions.
essential properties of the system can be captured Nodes in such networks generally represent
in an abstracted form with definite topological enzymes, with edges denoting interactions
structure which can be linked to functional prop- between them (Jeong et al. 2000).
erties of the system. This abstraction enables the • Immune networks: The mammalian immune
applications of tools developed in fields such as system can be considered as a number of
complex network theory, statistical physics, and interacting networks operating at different
sociology to be applied to biological networks, levels separated in both time and space.
greatly facilitating their analysis (Wuchty Within cells, gene networks link genes and
et al. 2003; Barabási and Oltvai 2004; Barabási transcription factors controlling gene expres-
and Albert 1999). It further enables the compar- sion; intracellular signaling networks link sur-
ison of biological systems to engineered systems face receptors on cells to gene-regulatory
which are traditionally described in network form events; other cells of the immune system
(e.g., as a flowchart) and thereby enables the such as B-cells and T-cells communicate via
identification of shared principles such as robust- mediators such as cytokines and hence form
ness, fragility, tolerance, and modularity (Alon cellular networks (Callard and Stark 2007).
2003). • Neuronal networks: The brain consists of intri-
Many biological systems have been studied cately organized networks of cortical connec-
from the perspective of network theory (Zhu tions; these connections can be modeled by
et al. 2007), for example: networks in which nodes are brain regions or
areas and edges represent fiber connections
• ▶ Transcription factors: Data obtained from between them (Kepes 2007).
chromatin immunoprecipitation (ChIP)
followed by probing of genomic microarrays The examples above provide a brief insight
and from ▶ DNA sequencing has been used to into the diversity of biological systems that can
assemble networks describing transcription be modeled using a network approach. Despite
factor binding sites in yeast and mammalian the obvious differences between the systems
cells. listed, the network approach provides a com-
• Protein-protein interactions: Such interac- mon framework in which these systems can be
tions represent the most readily available bio- analyzed and therefore compared. Such com-
logical networks to date. Data is generated parisons have revealed many similarities
from techniques such as two-hybrid assays between networks, not only those derived from
and other high-throughput techniques such as biological systems but also social systems (e.g.,
purification and mass spectrometry. In such friendship networks) and engineered systems
networks, nodes represent proteins with (e.g., the World Wide Web) (Barabási and
edges denoting physical interactions between Albert 1999). This tends to indicate generic
proteins (Jeong et al. 2001). Examples of par- principles of organization which can be
ticular protein interaction networks studied exploited to gain further understanding and
290 Biological Networks
will ultimately enable systems biology to com- tightly linked clusters. On the other hand, if
bine the numerous details about molecular C(k) follows C(k) k1, the network has a
interactions into a single framework, thereby hierarchical architecture meaning that
offering a means to address the structure of the sparsely connected nodes are part of highly
▶ cell as a whole. clustered regions with communication
between the different regions maintained by
a few hubs (Wuchty et al. 2001).
Basic Methodology
These properties can be used to classify the
Network topology plays a key role in understand- topology of a network into a number of models.
ing the architecture and function of biological Biological networks tend to be scale free in that
networks. A number of measures are commonly they are characterized by a degree distribution
used to describe the structure of a biological which follows a power law P(k) k g, i.e.,
network: most nodes in the network participate in very
few interactions, but a small number participate
• Degree: The degree of a node defines the in many and therefore function as hubs. Many
numbers of links the node has to other nodes biological systems are intrinsically modular,
and is the most basic measure used. The aver- i.e., functionality is achieved via a set of physi-
age degree <k> can be calculated; however, a cally or functionally linked modules which work
more useful measure is the degree distribution together. Modularity is accounted for in hierar-
P(k) which describes the number of nodes in chical network models which assume that clusters
the network having k links. A Poisson-shaped can combine together in an iterative manner to
degree distribution indicates that there are no generate a hierarchical or layered structure
highly connected nodes; in contrast, a power- (Barabási and Oltvai 2004).
law degree distribution indicates the presence
of a few highly connected nodes known
as hubs. Key Research Findings
• Mean shortest path: The shortest path is a
measure of the navigability of a network and The application of network analysis techniques to
describes the mean shortest distance between biological systems, made possible by the abstrac-
any two nodes in the network. Biological net- tion of biological systems as networks, indicates:
works (e.g., protein interaction networks and
transcriptional networks) exhibit a small- • Power-law degree distributions emerge as a
world property in which the mean shortest universal law characterizing cellular networks
path is very small. (Barabási and Albert 1999).
• Clustering coefficient: This measures the aver- • Many biological networks have the same
age probability in a network that two nodes topological scaling properties as complex
which have a mutual parent are also connected nonbiological systems such as social networks
(Watts and Strogatz 1998); using social net- or the World Wide Web, despite obvious dif-
work terminology, this can be interpreted as ferences in their components and structure,
“the friend of your friend is likely to be your and furthermore show striking similarities to
friend.” The clustering coefficient reveals the inherent organization of nonbiological
important information regarding network systems (Barabási and Albert 1999).
structure. C(k) measures the average cluster- • Integrity and robustness of biological net-
ing coefficient of nodes with k links; if C(k) is works are facilitated by structures which con-
independent of k, the network is either homo- tain a few highly connected hubs, e.g., genes
geneous or it is dominated by numerous small which can integrate multiple signals or trigger
Biological Networks 291
widespread attacks (Wuchty et al. 2003; Jeong roles. Identification of unknown pathogenic
et al. 2001); this ensures robustness against genes might help shed light on disease pathogenic
random failures but at the same time makes mechanisms (Zhu et al. 2007). Advancement in
them vulnerable to targeted attacks. This is our understanding of cytokine connectivity archi-
consistent with experimental data suggesting tectures has already demonstrated that immune B
that most mutations have little phenotypic cells do not function merely as individual clones
effect but that networks are extremely vulner- but work in innately integrated and hierarchical
able to attack at certain critical genes. collectives, shedding new light on immune-body
• The observed modularity found in biological systems organization and our understanding of
networks via network analysis supports the the role of the immune system in wound healing
notion that evolution acts over multiple levels and angiogenesis (Frankenstein et al. 2006).
(Wuchty et al. 2003); organisms can increase Manipulating the cytokine network has led to
in complexity via copying and reuse of novel approaches in treating acute and chronic
existing modules over time; furthermore, diseases, including cancer (Vilcek and Feldmann
modules can be readily reconfigured to adapt 2004).
to new conditions through evolution (Gerhart
and Kirschner 1997).
• Many different biological systems contain Future Directions
common motifs – subgraphs of a network
which are overrepresented when compared to Advancements in technology will increasingly
a randomized version of the same network. extend our abilities to accurately collect vast
For example, triangular motifs which have quantities of data. This will be coupled with
feed-forward functionality are found in both advancements in theoretical tools and methods
transcription-regulatory and feed-forward net- which will continue to drive forward understand-
works. Data obtained from yeast transcription ing of the complex interactions occurring within
networks points to the evolutionary conserva- cells, extending our understanding to
tion of key motif patterns with distinct intercellular networks and eventually extending
functionalities. it to cellular networks of complete organisms.
The science of complex networks has already
played a significant role in increasing understand-
Applications ing of the structural aspects of biological net-
works. Future research will additionally need to
Understanding the underlying structure of bio- take account of the dynamic aspects of cellular
logical networks may eventually enable a com- networks, including temporal and spatial aspects
plete description of the biological networks of a of activity and interactions within biological net-
complete cell to be elucidated, promising great works. Development of a novel framework in
benefits to medicine in the future. The identifica- which the function of biological networks can
tion of hubs – the highly connected nodes in a be interpreted heralds a new era of understanding
network – leads to the possibility of being able to of biology, disease pathologies, and medicine in
identify potential targets for drug design. The the future.
ability to target the high-degree proteins in a
network could lead, for example, to new strate-
gies for therapeutic mediation of signaling path- See Also
ways in cancer. Interactions in metabolic
networks are closely related to the ▶ gene func- ▶ Bioinformatics
tions and therefore have great potential for imme- ▶ Cell
diate applications in the interpretation of gene ▶ Evolution, Biological
292 Biological Radiation Effects
▶ Metabolism
▶ Protein Biological Safety Level
▶ Scale-Free Networks
Catharine A. Conley
Albert R, Jeong H, Barabási A-L (2000) Error and attack

tolerance in complex networks. Nature 406:378 Synonyms
Alon U (2003) The tinkerer as engineer. Science
301:1866–1867 BSL
Barabási A-L, Albert R (1999) Emergence of scaling in
random networks. Science 286:509–512
Barabási A-L, Oltvai Z (2004) Network biology: under-
standing the cells’ functional organization. Nat Rev Definition
Genet 5:101–113
Callard R, Stark J (2007) Networks of the immune system.
In: Kepes F (ed) Biological networks. Complex sys- The Biological Safety Level (BSL) is used in the
tems and interdisciplinary science. World-Scientific, USA to describe a set of specifications for the
Singapore precautions required to isolate potentially hazard-
Frankenstein Z, Alon U, Cohen I (2006) The immune- ous biological agents in a containment facility at a
body cytokine network defines a social architecture
of cell interactions. Biol Direct 1:32 specified stringency, ranging from BSL-1, which
Gerhart J, Kirschner MW (1997) Cells, embryos and evo- indicates the least strict containment, to BSL-4,
lution: towards a cellular and developmental under- which is the containment required for the most
standing of phenotypic variation and evolutionary hazardous human pathogens. A similar scale is
adaptability. Blackwell, Oxford
Jeong H, Tombor B, Albert R, Oltvai ZN, Barabási A-L used in the EU using the abbreviation P1-P4,
(2000) The large-scale organization of metabolic net- where P indicates either Pathogen or Protection.
works. Nature 407:651–654 For instance, a BSL-4 laboratory (P4 in Europe) is
Jeong H, Mason S, Barabási AL, Oltvai ZN (2001) Lethal- under reduced pressure relative to atmospheric
ity and centrality in protein networks. Nature
411:41–42 pressure, the exhausted air is filtered, and all
Kepes F (ed) (2007) Biological networks. Complex sys- wastes including water and any hardware leaving
tems and interdisciplinary science. World-Scientific, the high-containment area are sterilized chemi-
Singapore cally or by heating. Trained personnel work inside
Vilcek J, Feldmann N (2004) Historical review: cytokines
as therapeutics and targets of therapeutics. Trends the area in protective suits, overpressurized by air
Pharmacol Sci 25:201–209 pumped in from outside through a dedicated sys-
Watts DJ, Strogatz SH (1998) Collective dynamics of tem. The exterior of the suit is chemically
“small-world” networks. Nature 393(6684):409–410 disinfected when workers leave the area.
Wuchty S, Ravasz E, Barabaśi A-L (2003) The architec-
ture of biological networks. In: Deisboeck TS, Yasha
Kresh J, Kepler TB (eds) Complex systems in biomed-
icine. Kluwer, New York
Zhu X, Gerstein M, Snyder M (2007) Getting connected: Biological Sensors
analysis and principles of biological networks. Genes
Dev 21:1010–1024
▶ Biosensor
Biological Radiation Effects Biomarker, Isotopic
▶ Ionizing Radiation, Biological Effects ▶ Isotope Biosignatures

Biomarkers 293
Geochemists use “biomarkers” for ▶ molecular

Biomarkers fossils (fossil molecules such as steranes and
hopanes). Paleontologists use “biomarkers” for
Emmanuelle J. Javaux morphological and biosedimentary fossils.
Palaeobiogeology-Palaeobotany- Some scientists would use “biomarkers” for all B
Palaeopalynology, Geology Department, possible traces of life. The term “biosignatures”
Université de Liège, Liège, Belgium is very common in the literature as a general term
for all possible traces of life. However, astrobiol-
ogists now realize that discriminating possible
Keywords traces of life from unambiguous traces of life is
extremely difficult in the early Earth rock record
Chemical fossils; Molecular fossils; Morpholog- and even more challenging in an extraterrestrial
ical fossils context. Detecting life requires a set of multidis-
ciplinary approaches and criteria and a large (and
always improving but never completed) under-
standing of natural processes. Therefore, rather
Synonyms than using the term “signatures” or
“biosignatures,” the expression “traces of life”
Bioindice; Biosignature; Chemical fossil; Molec- or even better “indices of life” or “bioindices”
ular biomarkers; Molecular biosignatures; should be favored.
Morphological fossil; Trace of life
See Also
Definition
▶ Biogenicity
Biomarkers, biosignatures, and traces of life are ▶ Biomarkers, Morphological
three different terms or expressions related to the ▶ Biomarkers, Spectral
search for life in the Earth’s rock record or ▶ Biomineralization
beyond the Earth. However, they have different ▶ Dubiofossil
meanings, depending on the field of research ▶ Earth’s Atmosphere, History of the Origins
where they are used or depending on the “philos- ▶ Endogenicity
ophy” of the user. ▶ Fossil
▶ Molecular Fossils
▶ Pseudofossil
Overview ▶ Syngenicity
Biomarkers or “markers of life” or “traces of life”

include chemical, morphological, sedimentary,
or isotopic processes or structures that are bio- Botta O, Javaux EJ, Summons R, Rosing M, Bada J,
genic and could be detected to infer the past or Gomez Elvira J, Selsis F (eds) (2008) Strategies for
present presence of life. life detection ISSI space science series. Springer,
Heidelberg, Germany, p 380
However, in practice, astrophysicists use “bio-
Gargaud M, Mustin C, Reisse J (2009) Traces of past or
markers” for gases of possible biological origin present life: bio-signatures and potential life indica-
that could be detected in a planetary atmosphere. tors? Foreword. CR Palevol 8:593–603
294 Biomarkers Atmospheric, Evolution Over Geological Time
Biomarkers Atmospheric, Evolution Biomarkers, Morphological

Over Geological Time
Lisa Kaltenegger Palaeobiogeology-Palaeobotany-
Cornell University, Ithaca, NY, USA Palaeopalynology, Geology Department,
Definition
Keywords
The spectrum of the Earth has not been static
throughout the past 4.6 billion years. This is due Biominerals; Biosignatures; Fossils; Microfos-
to the temporal variations in atmospheric molec- sils; MISS; Stromatolites; Traces of life
ular abundances and temperature structure and in
the surface morphology and ▶ albedo. Observa-
tions of Earth-like planets at different geological Synonyms
ages will certainly help us understand the diver-
sity and some common features of terrestrial Biosignature; Trace of life
planets, including ours. For our own planet,
models show varying biomarkers over geological
times that could be detected in the spectrum (see, Definition
e.g., Schindler and Kasting 2000; Pavlov
et al. 2000; Kaltenegger et al. 2007) to character- Morphological biosignatures comprise objects or
ize it in terms of habitability and the possible structures of biological origin. They include body
presence of life. fossils, biominerals, and microbially influenced
sedimentary structures preserved in carbonates
(such as ▶ stromatolites), in siliciclastics (such
as MISS), or other rock types.
See Also
▶ Albedo Overview
▶ Biomarkers, Spectral
▶ Habitable Planet, Characterization Body fossils include microfossils and macrofos-
▶ Habitable Zone sils of various compositions and modes of fossil-
▶ Habitability of the Solar System ization. Microbially influenced sedimentary
structures include stromatolites (laminated car-
bonate rocks precipitated or trapped by microor-
ganisms), thrombolites (unlaminated carbonate
References and Further Reading rocks precipitated or trapped by microorgan-
isms), oncolites (small carbonate spheres with
Kaltenegger L, Traub WA, Jucks KW (2007) Spectral
evolution of an Earth-like planet. Astrophys
concentric laminations precipitated by microor-
J 658:598–616 ganisms), and MISS (a range of sedimentary
Pavlov AA, Kasting JF, Brown LL, Rages KA, Freedman structures produced by the presence and activity
R (2000) Greenhouse warming by CH4 in the atmo- of ▶ microbial mats in siliciclastic sediments).
sphere of early Earth. J Geophys Res 105:981–992
Schindler TL, Kasting JF (2000) Synthetic spectra of
Biominerals can be passively produced or
simulated terrestrial atmospheres containing possible actively produced by the organisms, and the
biomarker gases. Icarus 145:262–271 frontier between a biological or abiological
Biomarkers, Morphological 295
origin is tenuous. Ichnofossils or fossil traces of Key Research Findings and Applications
activities such as burrows, trails, and tracks are
sometimes considered as morphological bio- Body Fossils
markers. Abiotic processes can produce struc- A body fossil is any morphological remain of an
tures resembling microfossils (▶ Pseudofossil). organism after its death. It can have various sizes B
Chemical precipitates can look like stromato- (microscopic to macroscopic) and composition
lites. Minerals can self-assemble into complex (organic or mineral) and may represent a whole
structures that resemble body fossils or organism, part of an organism, or colonial organ-
biominerals. Deciphering the ▶ biogenicity of isms. Body fossils can have a variety of morphol-
an object or a structure is thus very difficult ogies and chemical composition, depending on
even using cutting-edge in situ ▶ techniques. their original properties and the conditions in
Several criteria have been proposed in the liter- which they are preserved. The organic composi-
ature in order to test their biogenicity. Most of tion may be transformed to various extents or
them underline the importance of proving the erased by the complex processes of fossilization.
endogenicity, the syngeneity, and the biogenicity The mineral composition may be primary
of the structures in question in a well- (produced or precipitated by the organism) or
characterized geological context. An additional secondary (the original carbonaceous or mineral
falsification approach is required to exclude all walls or envelopes being in that case partially or
possible abiotic hypotheses before a biological completely replaced by other minerals). Body
origin can be accepted. fossils may also be preserved as molds. In that
case, the fossils are first dissolved then the hole in
shape of the fossil is filled with a secondary
Basic Methodology mineral.
It is difficult to prove the biogenicity of micro-
Geobiological investigations in recent and past scopic body fossils in absence of carbonaceous
environments have determined the criteria for material. Microscopic cellular casts and molds
biogenicity of macroscopic and microscopic formed by various minerals can preserve or
morphological signatures of life. obscure the presence of a sheath and/or signifi-
Laboratory studies including artificial cell cantly alter the size of the microbes. For example,
degradation, bioalteration of minerals, and min- bacterial cells preserved by silicification
eralization experiments might reveal biotic pat- (replaced by silica) are difficult to discriminate
terns and preservable properties, although these from chemical precipitates. The organic compo-
studies may not reflect the full complexity of sition of a structure does not prove its biogenicity.
natural environmental conditions. Documenta- The term “organic” just indicates the presence of
tion of abiotic patterns and morphologies is also molecules with carbon and hydrogen. Organic
essential, but is difficult outside the lab on our matter is common in the universe and occurs in
biological planet. interstellar medium and in meteorites or is pro-
When trying to understand the origin of a duced abiotically by chemical reactions in hydro-
possible morphological biomarker, three main thermal conditions on Earth. Therefore, more
aspects need to be considered: (1) The criteria are required to assert the biogenicity of
preservational environments: what conditions carbonaceous material.
preserve cells with varying biochemical proper- Microscopic and microchemical analyses of
ties? (2) The taphonomy: how do processes of fossils isolated from shales or embedded in vari-
degradation and preservation retain, alter, or ous rocks are needed to demonstrate a range of
erase original biological properties? (3) The preservable biological properties, as well as to
criteria for biogenicity: how can we tell biologi- determine the taphonomic processes specific to
cal from nonbiological? the preservational environment and to the
296 Biomarkers, Morphological
original biology. The microfossils can be charac- variety of laminated or unlaminated precipitates
terized by their preserved walls (structure and and/or trapped and bound mineral grains. ▶ Stro-
ultrastructure) or sheaths, pigment and biopoly- matolites are found in the rock record since the
mer composition, morphology (regular cellular early Archean. They reach large sizes around
shape, complex shape, ornamentation, structured 2.7 Ga when their biogenicity is not questioned.
organic matter), abundance (population) and dis- Criteria such as rock fabrics, presence and distri-
tribution in the rock, orientation recording behav- bution of carbonaceous material or cells or fila-
ior and motility of benthic organisms (e.g., mats ments, unevenness of laminae, angle of repose of
with vertical filaments erected toward light), or laminae, etc., are investigated to prove or disprove
passive sedimentation of planktonic cells, pattern the biogenicity of these carbonate structures.
of cellular division, and, very importantly, their The process controlling the genesis of stro-
taphonomy (producing features such as collapsed matolites is not yet well understood, although
folded flexible hollow vesicles, sinuous seg- the association of macroscopic morphology with
mented filamentous shape, and pigmented sur- water energy is clear. Other structures indicating
faces of colonies). the presence of microbial mat interacting with the
sediment include particular fabric in sandstone,
Biominerals pinnacles, and other irregular mat surface mor-
Biominerals are minerals formed by organisms. phologies. Again, observations in natural settings
Biominerals can be passively produced or and experiments in the laboratory can help deter-
actively produced by the organisms, and the fron- mine criteria of biogenicity. The presence of
tier between a biological or abiological origin is body fossils strengthens the biogenicity of the
ambiguous. Biologically induced mineralization ▶ microbially induced sedimentary structures.
(BIM) includes biominerals formed through the
metabolic activity of microorganisms. Biologi-
cally controlled mineralization (BCM) includes Future Directions
biominerals whose formation is genetically con-
trolled by microorganisms. Associations of bio- Defining morphological (and other) biosignatures
logical tissues with minerals like bones or is not a simple task. The taphonomic processes
macroscopic and microscopic shells or plates differ depending on the preservational environ-
are easy to consider as body fossils and are exam- ments and the original biology, leading to bias in
ples of BCM. Microscopic mineral precipitates fossil diversity and morphology, often preventing
onto cells or sheaths or coated streamers or fila- identification or even recognition. A better under-
ment bundles are more difficult to interpret in standing of these environments and their particu-
absence of the cell or filament and can be BIM lar fossilization processes will help to make
or BCM. These precipitates may or may not form predictions of the types of biosignatures to search
in absence of the microorganisms. The contro- for in possible past or present extraterrestrial
versy regarding the magnetite minerals found in habitats (where and what to look for), important
the Martian meteorite ALH84001 is a good illus- when deciding landing sites and instrumentation
tration of the problem. Observations in natural for exobiological missions in situ and for returned
settings and experiments in the laboratory can samples. Characterizing the taphonomic pro-
help determine criteria for biogenicity. cesses and the criteria for recognition of
biogenicity will help interpret future observations
Microbially Induced Sedimentary Structures in early Earth and extraterrestrial records.
(MISS) and Stromatolites However, the early Archean record of stromat-
These structures are produced by the interaction olites and of microscopic biosignatures is still
between a microbial mat and the surrounding sed- actively debated, illustrating the difficulty
imentary environment. They can be preserved in (or impossibility) of defining unambiguous traces
carbonates or in siliciclastics. They include a of life in very old rocks, much less on another
Biomarkers, Spectral 297
planet where knowledge of the geological back- Grotzinger JP, Rothman DH (1996) An abiotic model for
ground is limited. Therefore, when searching for stromatolite morphogenesis. Nature 383:423–425
Hofmann HJ (2004) Archean microfossils and
traces of life in ancient and extraterrestrial mate- abiomorphs. Astrobiology 4(2):135–136
rials, a multitude of complementary studies and Knoll AH (2003) Life on a young planet. Princeton Uni-
biomarkers or bio-indices (not only morphologi- versity Press, Princeton, p 277 B
cal biomarkers) should be used. All possible abi- Konhauser KO (2007) Introduction to geomicrobiology.
Blackwell, Oxford, p 425
otic explanations should be considered and Krumbein WE, Paterson DM, Zvarzin GA (eds)
discarded before reaching an acceptable level of (2003) Fossil and recent biofilms. A natural history
reliability in any interpretation of biogenicity. of life on Earth. Springer, New York, p 504
Nevertheless, the conclusions will always be lim- Walter MR (ed) (1976) Developments in sedimentology,
vol 20, Stromatolites. Elsevier, New York
ited by our current understanding of the natural Westall F (1999) The nature of fossil bacteria: a guide to
processes involved. the search for extraterrestrial life. J Geophys Res
104:16437–16451
See Also
▶ Antarctica
Biomarkers, Spectral
▶ Biomarkers
▶ Biomineralization
Lisa Kaltenegger
▶ Dubiofossil
▶ Fossil
▶ Mars
Keywords
▶ Microbial Mats
▶ Microbially Induced Sedimentary Structures
Biomarkers; Extrasolar planets; Habitability;
▶ Microfossils, Analytical Techniques
Habitable zone; Planetary atmospheres;
▶ Pseudofossil
Spectroscopy
▶ Rio Tinto
▶ Stromatolites
Definition
The spectrum of a planet can contain signatures
Botta O, Javaux EJ, Summons R, Rosing M, Bada J, of atmospheric species that can remotely indicate
Gomez Elvira J, Selsis F (eds) (2008) Strategies for
habitable conditions. Some of the chemical spe-
life detection ISSI space science series. Springer,
Heidelberg, Germany, p 380 cies can indicate habitable conditions and some
Brasier MD, McLoughin N, Green O, Wacay D (2006) (combinations) can indicate biota on a planet. It is
A fresh look at the fossil evidence for early Archaean their presence and abundance along with other
cellular life. Philos Trans R Soc B 361:887–902
atmospheric species, in a certain context (e.g., the
Buick R (2001) Life in the Archean. In: Briggs DEG,
Crowther PR (eds) Paleobiology II. Blackwell Sci- properties of the star and the planet), that can
ence, London, pp 13–21 explore the underlying physics and characterize
Cady SL, Farmer JD, Grotzinger JP, Schopf JW, Steele a planetary environment. Here we concentrate on
A (2003) Morphological biosignatures and the search
characterizing a habitable planet using spectros-
for life on Mars. Astrobiology 3(2):351–369
Fortin F (2004) What biogenic minerals tell us. Science copy and discuss which features could point
303:1618–1619 toward a biological origin (see also ▶ Bio-
Gargaud M, Mustin C, Reisse J, Vandenabeele- markers, atmospheric; ▶ Habitable planet, char-
Trambouze O (eds) (2009) Traces de vie présente ou
acterization; Habitability, effects of eccentricity;
passée: quels indices, signatures ou marqueurs?
Comptes rendus Académie des Sciences Paris, ▶ Habitability, effects of stellar irradiation;
PalEvol 8, 691 p ▶ Habitability of the solar system).
298 Biomarkers, Spectral
History of 3 for rocky planets that could potentially sup-

port life. Searches will concentrate on planets in
Sagan et al. (1993) analyzed a spectrum of the the so-called ▶ habitable zone (HZ) where water
Earth taken by the Galileo probe, searching for could remain liquid if present and gases produced
signatures of life, and concluded that the large by biota could easily exchange with and influence
amount of O2 and the simultaneous presence of the atmosphere. The HZ is defined for surface
traces of CH4 are strongly suggestive of biology. conditions only.
After a decade rich in giant exoplanet detections, Our search for signs of life is based on the
observational techniques have now reached the assumption that extraterrestrial life shares funda-
ability to find planets of less than 10 MEarth mental characteristics with life on Earth, in that it
(so-called ▶ Super-Earths) that may potentially requires liquid water as a solvent and has a
be habitable. carbon-based chemistry (see, e.g., Brack 1993;
Des Marais et al. 2002). Life on the basis of a
different chemistry is not considered here,
Overview because the vast range of conceivably possible
life-forms might produce signatures in their
We discuss how we can read a planet’s spectrum atmosphere that are so far unknown. Therefore,
to assess its habitability and search for the spec- we assume that extraterrestrial life is similar to
tral signatures of a biosphere. To characterize a life on Earth in its use of the same input and
planet’s atmosphere and its potential habitability, output gases and that it exists out of thermody-
we look for absorption features in the reflection namic equilibrium (Lovelock 1975). “Bio-
and transmission spectrum of the planet. On markers” here refer to detectable species or a set
Earth, some atmospheric species exhibiting of species, whose presence at significant abun-
noticeable spectral features in the planet’s spec- dance strongly suggests a biological origin (e.g.,
trum result directly or indirectly from biological couple CH4 + O2 or CH4 + O3 (Lovelock 1975)).
activity: The main ones are O2, O3, CH4, and A bioindicator is maybe indicative of biological
N2O. CO2 and H2O are in addition important as processes but can also be produced abiotically.
greenhouse gases in a planet’s atmosphere and Chemoautotrophic life, whose metabolism
potential sources for high O2 concentration from does not depend on the stellar light, can still exist
photosynthesis. Future remote-sensing character- outside the HZ, thriving in the interior of the planet
ization of planetary environments can be used to where liquid water is available. Such metabolisms
test our understanding of the factors that contrib- (see Extremophile) rely on very limited sources of
ute to planetary habitability. To detect such fea- energy (compared to stellar light) and electron
tures remotely with first-generation spectroscopy donors (compared to H2O on Earth). They mainly
mission that will not resolve the planet, the atmo- catalyze reactions that would occur at a slower rate
sphere has to show such chemical features in purely abiotic conditions, and they are thus not
globally. expected to modify a whole planetary environ-
ment in a way detectable remotely.
Introduction
Dedicated future space missions will have the
explicit purpose of detecting other Earth-like Basic Methodology
worlds, analyzing their characteristics, determin-
ing the composition of their atmospheres, inves- Atmospheric Features of a Habitable Planet
tigating their capability to sustain life as we know A planet observed remotely is a very faint, small
it, and searching for signs of life. This can set our object close to a very bright and large object, its
own planet, the only known habitat, in context parent star. The Earth is about a million times
with other rocky worlds and expand our statistics fainter than the Sun in the mid-infrared and about
a billion times fainter than the Sun in the visible. conditions and, possibly, the presence of a bio-
Suppressing the starlight to that degree allows sphere: CO2, H2O, O3, CH4, and N2O in the
collecting the planet’s light and characterizing thermal infrared and H2O, O3, O2, CH4, and
its atmosphere. CO2 in the visible to near infrared in reflection
(Fig. 1) and transmission spectra (Fig. 2). The B
presence or absence of these spectral features
Key Research Findings (detected individually or collectively) will indi-
cate similarities or differences among the atmo-
Spectra of Earth in Reflection, Emission, and spheres of terrestrial planets and their
Transmission astrobiological potential.
Figure 1 shows observations and model fits to the
spectra of the Earth in three wavelength ranges, Characterizing Planetary Environments
the visible and near infrared represent the It is relatively straightforward to remotely ascer-
reflected starlight, while the infrared tain that the Earth is a habitable planet, replete
(IR) spectrum is generated by the emitted heat with oceans, a greenhouse atmosphere, global geo-
flux of the planet (Kaltenegger et al. 2007). For chemical cycles, and life – if one has data with
the data shown in Fig. 1 (a) is the visible earth- arbitrarily high signal-to-noise ratio and spatial
shine spectrum (Woolf et al. 2002), (b) is the and spectral resolutions. The interpretation of
near-infrared earthshine (defined as light from observations of other planets with limited signal-
the Sun reflected from the Earth to the Moon to-noise ratio and spectral resolution, as well as
and back again) spectrum (Turnbull et al. 2006), absolutely no spatial resolution, as envisioned for
and (c) is the thermal infrared spectrum of the the first-generation instruments, will be far more
Earth as measured by a spectrometer en route to challenging. This implies that we need to gather
Mars (Christensen and Pearl 1997). The data are information on the planet’s environment to under-
shown in black and the Smithsonian Astrophysi- stand what we will see. We can then test if we have
cal Observatory model in red. In each case, the an abiotic explanation of all compounds seen in
constituent gas spectra in a clear atmosphere are the atmosphere of such a planet. If we do not, we
shown below, illustrating how each molecule can work with the exciting biotic hypothesis.
contributes to the overall spectrum. O2 and O3 in combination with a reducing gas
Figure 2a shows observations (Irione 2002) like CH4 are good biomarker candidates.
and model fits to transmission spectra of the Reduced gases and oxygen have to be produced
Earth (Kaltenegger and Traub 2009) that repre- concurrently to be detectable in the atmosphere,
sent the transit of the Earth. The data are shown in as they react rapidly with each other. Thus, the
blue and the model in red. A transmission spec- chemical imbalance traced by the simultaneous
trum gives the apparent radius of a planet versus signature of O2 and/or O3 and of a reduced gas
wavelength. Different molecules in the planet’s like CH4 can be considered as a signature of
atmosphere absorb part of the stellar light and biological activity (Lovelock 1975). These spe-
make the planet appear bigger than its solid cies can be detected by a low-resolution spectro-
radius at characteristic wavelengths, depending graph (O2 in the visible with a resolution <100,
on the molecule. This is shown here as effective O3 and CH4 in the infrared with a resolution
height in the atmosphere. The atmospheric fea- <50). Note that if the presence of biogenic
tures are indicated. For details on the Earth’s gases such as O2/O3 + CH4 may imply the pres-
transmission spectrum during a lunar eclipse, ence of a massive and active biosphere, their
see Palle et al. (2009) and Vidal-Madjar absence does not imply the absence of life. Life
et al. (2010). existed on Earth before the interplay between
Both spectral regions contain the signature of oxygenic photosynthesis and carbon cycling pro-
atmospheric gases that may indicate habitable duced an oxygen-rich atmosphere.
a 1.2
Data
1
Model
0.8
Relative reflectance 0.6
0.4
1
0.8 O3 O2 O2 O2 H2O
0.6
0.4
0.2
0
0.6 0.8
Wavelength (mm)
b c
0.8
10
Data
0.6 Model
Data
0.4 Model 5
0.2
0
0 1
0.8 CH4 CH4
H2O O2 CO2
0.6 H2O O3 CO2 H2O
0.4
0.2
0 0
1 1.2 1.4 1.6 1.8 5 10 15 20
Wavelength (mm) Wavelength (mm)
Biomarkers, Spectral, Fig. 1 Observed reflectivity integrated Earth, as determined from earthshine and
spectrum in the visible (Woolf et al. 2002) (a) and near space, respectively. The data are shown in black and the
infrared (Turnbull et al. 2006) (b) and emission spectrum Smithsonian Astrophysical Observatory model in red. The
in the infrared (IR) (Christensen and Pearl 1997) (c) of the reflectivity scale is arbitrary
Temperature and Radius of a Planet detailed information on the thermal profile of a

Knowing the temperature and planetary radius is planetary atmosphere (see atmospheric struc-
crucial for a general understanding of the physi- ture). This, however, requires a spectral resolu-
cal and chemical processes occurring on the tion and a sensitivity that are well beyond the
planet (e.g., tectonism, ▶ atmosphere, escape). performance of a first-generation spacecraft.
In theory, spectroscopy can provide some Generally, the surface temperature of a planet at
Biomarkers, Spectral, a 1
Fig. 2 Synthetic
90 km
transmission spectra of the 10
Earth from UV to IR
75 km
(Kaltenegger and Traub 10
2009) for a cloudless
50 km B
atmosphere based on 10
shuttle data (Irione 2002) 40 km
and calculated for a line of 10
(Relative transmission)
sight at the tangent height 30 km
indicated on the right (a). 10
A transmission spectrum 25 km
gives the apparent radius of 10
a planet versus wavelength. 20 km
Different molecules in the 10
planet’s atmosphere absorb 12 km
the stellar light and make 10
the planet appear bigger 10 km
than its solid radius at 10
characteristic wavelengths, 6 km
10
depending on the molecule.
This is shown here as 4 km
10
effective height in the
atmosphere (b). The 1 km
0
atmospheric features are 8 10 12 14 16
indicated below Wavelength (mm)
b
60
CO2
Effective height
CO2
H2O O3
40 O
3
O2 H2O CH4
20 HNO3 H2O
0
5 10 15 20
1
0.8
O2 HNO3 H2O
0.6 H2O CH
4
O3 H2O O3
0.4 CO2
Rel. transmission
CO2
0.2
0
5 10 15 20
1
0.8
HNO3 H2O
0.6 O2 CH4
O3
0.4
CO2
0.2
0
5 10 15 20
Wavelength (mm)
a specific distance from its star depends on its photosynthetic bacteria (cyanobacteria) to use
▶ albedo and on the greenhouse warming by abundant H2O, instead of having to rely on scarce
atmospheric compounds. However, even with a supplies of electron donors like H2 and H2S to
low-resolution spectrum of the thermal emission, reduce CO2. With oxygenic photosynthesis, the
the mean effective temperature can be obtained production of the biomass becomes limited only
from the peak of the blackbody curve in the by nutrients and no longer by energy (light in this
infrared (IR). The measured IR flux can directly case) nor by the abundance of electron donors.
be converted into a brightness temperature that Oxygenic photosynthesis at a planetary scale
will provide information on the temperature of results in the storage of large amounts of radiative
the atmospheric layers responsible for the emis- energy in chemical energy, in the form of organic
sion. The overall flux of the planet in the IR is matter. For this reason, oxygenic photosynthesis
determined by the surface area of the planet and had a tremendous impact on biogeochemical
its effective temperature; therefore, the radius of cycles on Earth and eventually resulted in the
the emitting surface (e.g., ground or cloud layers; global transformation of the Earth’s environ-
see ▶ Clouds) of the planet can be obtained. ment. Less than 1 ppm of atmospheric O2 comes
The ability to associate the spectrum with a from abiotic processes (Walker 1977).
surface temperature relies on the existence and Cyanobacteria and plants are responsible for the
identification of spectral windows probing the biotic production by using solar photons to
surface or specific atmospheric levels. Such iden- extract hydrogen from water and using it to pro-
tification is not trivial. For an Earth-like planet, duce organic molecules from CO2. This metabo-
there are some atmospheric windows that can be lism is called oxygenic photosynthesis. The
used in most of the cases, especially between reverse reaction, using O2 to oxidize the organics
8 and 11 mm. This window would, however, produced by photosynthesis, can occur abioti-
become opaque at high H2O partial pressure cally when organics are exposed to free oxygen
(e.g., in the inner part of the habitable zone or biotically by both eukaryotes and certain pro-
(HZ) where a lot of water is vaporized) and at karyotes using O2 and consuming organics.
high CO2 pressure (e.g., a very young Earth or the Due to a balance in the above processes, the
outer part of the HZ). The accuracy of the radius net release of O2 in the atmosphere actually
and temperature determination will depend on depends on the burial of organics in sediments.
the quality of the fit (and thus on the sensitivity Each reduced carbon buried results in a free O2
and resolution of the spectrum), the precision of molecule in the atmosphere. This net release rate
the Sun-star distance, the cloud coverage, and is also counterbalanced by the weathering of fos-
also the distribution of brightness temperatures silized carbon when exposed to the surface. The
over the planetary surface. Assuming the effec- oxidation of reduced volcanic gases such as H2
tive temperature of our planet were radiated from and H2S also accounts for a significant fraction of
the uppermost cloud deck at about 12 km would the oxygen losses. The atmospheric oxygen is
introduce about 2 % error on the Earth’s radius recycled through respiration and photosynthesis
derived from emergent and/or transmission spec- in less than 10,000 years. In the case of a total
tra. For transiting planets, the accuracy of the extinction of the Earth’s biosphere, the atmo-
radius of the planet depends on how well the spheric O2 would disappear in a few million
host star is characterized. years.
The spectrum of the Earth has exhibited a
Potential Biomarkers strong infrared signature of ozone for more than
Owen (1980) suggested searching for O2 as a 2 billion years and a strong visible signature of O2
tracer of life. Oxygen in high abundance is a for an undetermined period of time between 2 and
promising bioindicator. Oxygenic photosynthe- 0.8 billion years (depending on the required depth
sis, the by-product of which is molecular oxygen of the band for detection and also the actual
extracted from water, allows terrestrial plants and evolution of the O2 level; Kaltenegger
et al. 2007). This difference is due to the fact that the Earth’s atmosphere by the biosphere with
a saturated ozone band appears already at very broadly similar rates, but the atmospheric abun-
low levels of O2 (104 ppm), while the oxygen dance of NH3 is orders of magnitude lower due to
line remains unsaturated at values below the pre- its very short lifetime under UV irradiation. The
sent atmospheric level (Segura et al. 2003). The detection of NH3 in the atmosphere of a habitable B
depth of the saturated O3 band is determined by planet would thus be extremely interesting, espe-
the temperature difference between the surface- cially if found with oxidized species. The detec-
to-clouds region (producing the spectral contin- tion of H2O and CO2 are important in the search
uum) and the ozone layer. for signs of life not as biosignatures themselves
N2O is produced in abundance by microbial but because they are raw materials for life and
life but only in negligible amounts by abiotic thus necessary for planetary habitability.
processes. Nearly all of Earth’s N2O is produced There are other molecules that could, under
by the activities of anaerobic nitrifying and some circumstances, act as excellent biomarkers;
denitrifying bacteria. N2O would nevertheless for example, the manufactured chlorofluorocar-
be hard to detect in the Earth’s atmosphere with bons (CCl2F2 and CCl3F) are observed in our
low resolution, as its abundance is low at the current atmosphere in the thermal infrared wave
surface (0.3 ppmv) and falls off rapidly in the band, but their abundances are currently too low
stratosphere. Spectral features of N2O would to be spectroscopically observed at low
become more apparent in atmospheres with resolution.
more N2O and/or less H2O vapor. Segura
et al. (2003) have calculated the level of N2O
for different O2 levels and found that its abun- Applications
dance decreases together with O2.
The methane found in the present atmosphere Low-Resolution Spectral Information in the
of the Earth has a biological origin, except for a Visible to Near IR
small fraction produced abiotically in hydrother- In the visible to near infrared, one can see
mal systems where hydrogen is released by the increasingly strong telluric H2O bands at 0.73,
oxidation of ferrous iron by H2O and reacts with 0.82, 0.95, and 1.14 mm. The strongest O2 feature
CO2. Depending on the degree of oxidation of a is the saturated Fraunhofer A-band at 0.76 mm.
planet’s crust and upper mantle, such A weaker feature at 0.69 mm cannot be seen with
nonbiological mechanisms can produce large low resolution. O3 has a broad feature, the
amounts of CH4 under certain circumstances. Chappuis band, which appears as a broad trian-
Therefore, the detection of methane alone cannot gular dip in the middle of the visible spectrum
be considered a sign of life, while its detection in from about 0.45 to 0.74 mm. The feature is very
an oxygen-rich atmosphere would be difficult to broad and shallow. Methane at present terrestrial
explain in the absence of a biosphere. Note that abundance (1.65 ppm) has no significant visible
methane on Mars has been suggested (Mumma absorption features, but at high abundance it has
et al. 2009), while the atmosphere of Mars con- strong visible bands at 0.88 and 1.04 mm, readily
tains 0.1 % of O2 and some ozone. In this case, the detectable, for example, in early Earth models.
amounts involved are extremely low, and the CO2 has negligible visible features at present
origin of the Martian O2 and O3 is known to be Earth abundance, but in a high CO2 atmosphere
photochemical reactions initiated by the photol- of 10 % CO2, like in an early Earth evolution
ysis of CO2 and water vapor. If confirmed, the stage, the weak 1.06-mm band could be observed.
presence of methane could be explained by sub- In the UV, O3 shows a strong feature that is not
surface geochemical process, assuming that discussed here. The red edge of land plants due to
reducing conditions exist on Mars below the chlorophyll developed about 0.44 Ga. It could be
highly oxidized surface. The case of NH3 is sim- observed on a cloudless Earth or if the cloud
ilar to the one of CH4. They are both released into pattern is known.
Low-Resolution Spectral Information in the steady state maintained by photodissociation by

Mid-IR the stellar UV radiation but with a constant ele-
In the mid-IR for Earth, the detectable signatures mental composition of the atmosphere, while the
of biological activity in low resolution are the second one is a net source of oxygen. In order to
combined detection of the 9.6-mm O3 band, the reach detectable levels of O2 (in the reflected
15-mm CO2 band, and the 6.3-mm H2O band or its spectrum), the photolysis of CO2 has to occur in
rotational band that extends from 12 mm out into the absence of outgassing of reduced species and
the microwave region (Selsis 2003). The 9.6-mm in the absence of liquid water, because of the wet
O3 band is highly saturated and is thus a poor deposition of oxidized species. Normally, the
quantitative indicator but an excellent qualitative detection of the water vapor bands simulta-
indicator for the existence of even traces of O2. neously with the O2 band can rule out this abiotic
CH4 is not readily identified using low-resolution mechanism (Segura et al. 2007), although one
spectroscopy for present-day Earth, but the meth- should be careful, as the vapor pressure of H2O
ane feature at 7.66 mm in the IR is easily detect- over a high-albedo icy surface might be high
able at higher abundances (e.g., 100 on early enough to produce detectable H2O bands. In the
Earth; Kaltenegger et al. 2007), provided that the infrared, this process cannot produce a detectable
spectrum contains the whole band and a high O3 feature (Selsis et al. 2002). The loss of hydro-
enough signal-to-noise ratio. Taken together gen to space can result in massive oxygen left-
with molecular oxygen, abundant CH4 can indi- overs: More than 200 bars of oxygen could build
cate biological processes (see also Sagan up after the loss of the hydrogen contained in the
et al. 1993; Segura et al. 2003). Although meth- Earth’s oceans. However, the case of Venus tells
ane’s abundance is less than 2 ppm in the Earth’s us that such oxygen leftover has a limited lifetime
atmosphere, the 7.75-mm band shows up in a in the atmosphere (because of the oxidation of the
medium-resolution (Res = 100) infrared spec- crust and the loss of oxygen to space): We do not
trum. Three N2O features in the thermal infrared find O2 in the Venusian atmosphere despite the
are detectable at 7.75 and 8.52 mm and at massive loss of water probably experienced in the
16.89 mm for levels higher than in the present early history of the planet. Also, such
atmosphere of the Earth. evaporation-induced buildup of O2 should occur
only closer to a certain distance from the star and
Abiotic Sources of Biomarkers affect small planets with low gravity more dra-
Abiotic sources of biomarkers are very important matically than more massive planets. For small
to assess, so that we can identify when they might planets (<0.5 MEarth) close to the inner edge of
constitute a “▶ false positive” for life. CH4 is an the habitable zone, there is a risk of abiotic oxy-
abundant constituent of the cold planetary atmo- gen detection, but this risk becomes negligible for
spheres in the outer solar system. On Earth, it is big planets further away from their star. The fact
produced abiotically in hydrothermal systems that on the Earth oxygen and indirectly ozone are
where H2 (produced from the oxidation of Fe by by-products of the biological activity does not
water) reacts with CO2 in a certain range of mean that life is the only process able to enrich
pressures and temperatures. In the absence of an atmosphere with these compounds. The ques-
atmospheric oxygen, abiotic methane could tion of the abiotic synthesis of biomarkers is
build up to detectable levels. Therefore, the crucial, but only very few studies have been ded-
detection of CH4 cannot be attributed unambigu- icated to it (Rosenqvist and Chassefiere 1995;
ously to life. Leger et al. 1993; Lagrange et al. 2009; Selsis
O2 also has abiotic sources, e.g., (a) the pho- et al. 2002).
tolysis of CO2, followed by recombination of
O atoms to form O2 (O + O + M ! O2 + M), Cryptic Worlds
and (b) the photolysis of H2O followed by escape On the Earth, photosynthetic organisms are
of hydrogen to space. The first source results in a responsible for the production of nearly all of
the oxygen in the atmosphere. However, in many temperature, and biota) expand our knowledge
regions of the Earth, and particularly where sur- of how a habitable planet’s spectrum could
face conditions are extreme, for example, in hot appear and how robust our definition of bio-
and cold deserts, photosynthetic organisms can markers is. We are just starting to learn how to
be driven into and under substrates where light is characterize habitable planets remotely. The B
still sufficient for photosynthesis. These commu- results of a first-generation mission will most
nities exhibit no detectable surface spectral sig- likely result in an amazing scope of diverse
nature. The same is true of the assemblages of planets that will place our own planet in an over-
photosynthetic organisms at more than a few all context.
meters depth in water bodies. These communities
are widespread and dominate local photosyn-
thetic productivity on Earth. Such worlds could See Also
be very interesting Earth analogues that could be
habitats but cannot exhibit a biological surface ▶ Albedo
feature, such as that due to chlorophyll, in the ▶ Atmosphere, Structure
disk-averaged spectrum (Cockell et al. 2009). ▶ Biomarkers Atmospheric, Evolution Over
Geological Time
▶ Clouds
Summary ▶ Greenhouse Effect
▶ Habitability, Effect of Eccentricity
Dedicated future space missions will have the ▶ Habitability, Effects of Stellar Irradiation
explicit purpose of detecting other Earth-like ▶ Habitability of the Solar System
worlds, analyzing their characteristics, determin- ▶ Habitable Planet, Characterization
ing the composition of their atmospheres, inves- ▶ Habitable Zone
tigating their capability to sustain life as we know ▶ Habitable Zone, Effect of Tidal Locking
it, and searching for signs of life. These searches ▶ Mars
for signs of life will be based on the assumption ▶ Super-Earths
that extraterrestrial life shares fundamental char- ▶ Venus
acteristics with life on Earth, in that it requires
liquid water as a solvent, has an energy source,
and has a carbon-based chemistry. Life on the References and Further Reading
basis of a different chemistry is not considered
here, because the vast possible range of conceiv- Arnold L, Gillet S, Lardiere O, Riaud P, Schneider
J (2002) A test for the search for life on extrasolar
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identified some remotely detectable biomarkers 392:231–237
whose presence at significant abundance would Brack A (1993) Liquid water and the origin of life. Orig
Life Evol Biosph 23(1):3–10
strongly suggest a biological origin (e.g., the cou- Christensen PR, Pearl JC (1997) Initial data from the Mars
ples CH4 + O2 or CH4 + O3 (Lovelock 1975) or global surveyor thermal emission spectrometer exper-
N2O). In addition to searching for signs of life, we iment: observations of the Earth. J Geophys Res
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Cockell CS, Kaltenegger L, Raven JA (2009) Cryptic
physically and chemically works. photosynthesis – extrasolar planetary oxygen without
a surface biological signature. Astrobiology
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Biologically controlled biomineralization; Bio-
Segura A, Krelove K, Kasting JF, Sommerlatt D,
Meadows V, Crisp D, Cohen M, Mlawer E (2003) logically induced biomineralization;
Ozone concentrations and ultraviolet fluxes on Biosignature; Bioprecipitate
Biomineralization 307
Synonyms with particular shapes. Alternatively, nuclei

may aggregate, forming mesocrystals, some-
Bioprecipitation times with the mediation of an organic matrix
(e.g., Sethmann et al. 2006). This mechanism
has been increasingly identified in B
Definition biomineralizing systems, but exact molecular
mechanisms remain unknown.
Biomineralization is the process by which organ-
isms form minerals, which are called Three types of biomineralization processes
biominerals. It is a widespread phenomenon have been distinguished in the literature
induced or controlled by a huge diversity of (Dupraz et al. 2009):
eukaryotes, bacteria, or archaea. There is a very
large diversity of chemical compositions and 1. Biologically controlled biomineralization
structures for biominerals, including carbonates refers to cases in which a specific cellular
(e.g., aragonite, calcite), silicates (e.g., opal), and activity directs the nucleation, growth, mor-
Fe and/or Mn oxides. Biominerals can be used in phology, and final location of a mineral,
the search for traces of ancient terrestrial or extra- involving specific genes. Many eukaryote
terrestrial life. They can be used as recorders of biominerals result from this process. Forma-
paleoenvironmental conditions, especially in tion of intracellular magnetites in
paleoclimatology (e.g., corals, foraminifers). magnetotactic bacteria is another emblematic
Finally, their unique properties can inspire example.
mimetic strategies intending to design 2. Biologically induced biomineralization results
nanomaterials for new advanced technologies. from the indirect modification of the chemis-
try of the environment by biological activity.
Several types of microbial metabolisms can,
Overview for example, modify solution chemistry and
induce carbonatation, i.e., precipitation of car-
Organisms can impact mineral formation at sev- bonate minerals, including photosynthesis,
eral steps: sulfate reduction, urea degradation, ammoni-
fication, or denitrification (Castanier
1. Organisms can modify actively or passively et al. 2000).
solution chemistry in their vicinity, by chang- 3. Biologically influenced biomineralization is
ing pH or the activity of any chemical species. defined as passive mineralization of organic
This may raise supersaturation with a mineral matter, whose properties influence crystal
phase in the surrounding solution and makes morphology and composition. Polysaccha-
conditions thermodynamically favorable to rides and proteins have been shown to impact
mineral precipitation. biomineralization in many cases (e.g., Obst
2. Organisms can impact nucleation, i.e., forma- et al. 2009). The term organomineralization
tion of very small and unstable mineral seeds. is sometimes used, encompassing biologically
They can indeed stabilize these nuclei by pro- influenced and biologically induced
viding large surfaces (e.g., cell walls, extra- biomineralization.
cellular polymers) with lower surface tension.
3. Finally, organisms can impact mineral The morphology, the structure, and the chem-
growth. Traditionally, growth takes place istry of prokaryote biominerals have frequently
layer by layer. Organisms can inhibit crystal been proposed as potential biosignatures (e.g.,
growth along certain directions by production Konhauser 1998) and used to infer the presence
of poisoning molecules, forming minerals of traces of life in ancient terrestrial or
308 Biopan
extraterrestrial rocks such as the Martian meteor-

ite ALH84001 (McKay et al. 1996). For further Biopan
reading, reviews on various biomineralizing sys-
tems can be found (e.g., Bäuerlein 2000; Mann René Demets
2001; Weiner and Dove 2003; Benzerara and ESTEC (HSF-USL), Noordwijk, The Netherlands
Menguy 2009).
Keywords
See Also Exposure experiments; Radiation biology
▶ Biomarkers, Morphological
▶ Bioprecipitation Definition
▶ Mineral
Biopan is a pan-shaped exposure facility for
experiments in the domains of astrobiology, radi-
ation biology, and radiation dosimetry (Demets
References and Further Reading et al. 2005). Externally mounted on unmanned
Bäuerlein E (2000) Biomineralization: from biology to
recoverable satellites of the Foton type, Biopan
biotechnology and medical application. Wiley-VCH, flies 2-week missions in low Earth orbit at 63.0
Weinheim inclination, allowing exposure of biological sam-
Benzerara K, Menguy N (2009) Looking for traces of life ples to the harsh space conditions. Six flights
in minerals. CR Palevol 8:617–628
have been completed between 1992 and 2007
Castanier S, Le Métayer-Levrel G, Perthuisot JP
(2000) Bacterial roles in the precipitation of carbonate with up to ten different experiments per flight.
minerals. In: Riding R, Awramik SM (eds) Microbial Biopan carries its experiment packages (total
sediments. Springer, Heidelberg, pp 32–39 mass 4 kg max.) on two mounting plates (total
Dupraz C, Reid RP, Braissant O, Decho AW, Norman SR,
surface area 1,080 cm2).
Visscher PT (2009) Processes of carbonate precipita-
tion in modern microbial mats. Earth Sci Rev
96:141–162
Konhauser KO (1998) Diversity of bacterial iron mineral- Overview
ization. Earth Sci Rev 43:91–121
Mann S (2001) Biomineralization: principles and concepts
in bioinorganic materials chemistry. Oxford Univer- In orbit, the hinged lid of Biopan is opened by
sity Press, Oxford telecommand whereupon the experiments are
McKay DS, Gibson EK, Thomas-Keprta KL, Vali H, freely exposed to the space environment. At the
Romanek CS, Clemett SJ, Chillier XDF, Maechling
end of the flight, the lid is hermetically closed and
CR, Zare RN (1996) Search for past life on Mars:
possible relic biogenic activity in Martian meteorite locked. During reentry into the atmosphere,
ALH84001. Science 273:924–930 Biopan and its contents are protected against the
Obst M, Wehrli B, Dittrich M (2009) CaCO3 nucleation by frictional heat by an ablative heat shield. Biopan
cyanobacteria: laboratory evidence for a passive,
is equipped with a variety of sensors to monitor
surface-induced mechanism. Geobiology 7:324–347
Sethmann I, Hinrichs R, Worheide G, Putnis A (2006) and record the environmental history of the test
Nano-cluster composite structure of calcitic sponge samples. Included are ultraviolet (UV) sensors, a
spicules – a case study of basic characteristics of radiometer, and a set of eight thermistors to mea-
biominerals. J Inorg Biochem 100:88–96
sure the experiment temperatures. The sensor
Weiner S, Dove PM (2003) An overview of biominerali-
zation processes and the problem of the vital effect. data are stored on board and retrieved after land-
Rev Mineral Geochem 54:1–29 ing. The temperature profile of the experiments is
Biopolymer 309
selectable. A noncontrolled mode can be chosen

with temperatures freely oscillating between Biopolymer
20 C and +10 C, in sync with the alter-
nating periods of solar illumination and Lucas J. Stal
shadowing in orbit. Alternatively, by using elec- Department of Marine Microbiology, Royal B
trical heaters and thermal blankets, a stable tem- Netherlands Institute of Sea Research (NIOZ),
perature can be provided with a fixed set point in Yerseke, The Netherlands
the 10–25 C range. Organic molecules, bacterial
spores, ▶ archaea, plant seeds, ▶ lichens, and
tardigrades have been exposed in Biopan to a Keywords
combination of solar UV, space vacuum, space
radiation, wide temperature fluctuations, and Biopolymer; Cellulose; Chitin; Exopolymers;
weightlessness. Biopan carries reference sam- Heteropolymer; Homopolymer; Nucleic acid;
ples, which are kept under identical conditions Peptidoglycan; Polyhydroxyalkanoate; Polypep-
but shielded against UV. Additional control sam- tide; Polyphosphate; Polysaccharide; Protein;
ples are maintained on ground. The typical oper- Silicate; Starch
ational cycle of Biopan includes experiment
integration at 1 week before launch, 2 weeks of
orbital flight, and return of the experiments to the Synonyms
investigators at 4 days after landing. Biopan was
designed and built for ESA by Kayser-Threde Natural polymer
(Munich, Germany) with Kayser Italia (Livorno,
Italy) responsible for the flight software and the
electronics. The heat shield is manufactured by Definition
TsSKB-Progress (Samara, Russia).
Biopolymers are mostly organic molecules that
are composed of repeating monomers and pro-
duced by living organisms. Homopolymers are
See Also such molecules that are composed of one
type of monomer, while heteropolymers are com-
▶ Exposure Facilities posed of more than one type of monomer. Such
▶ Foton Capsule, Spacecraft ▶ polymers may either be a repeating unit of two
or more monomers or may be more
heterogeneous, complex, and often branched.

Overview
Demets, R., Schulte, W., & Baglioni, P. (2005). The past,
present and future of BIOPAN. Adv Space Res, 36(2),
311–316. There is a wealth of examples of biopolymers
important for the organisms that produce them
and for the ecosystems they inhabit. Polynucleo-
tides and polypeptides are the key to life. Poly-
Biopoesis nucleotides (deoxyribonucleic acid, DNA, and
ribonucleic acid, RNA) are the carriers of infor-
▶ Origin of Life mation in cells and are responsible for the
310 Bioprecipitation
translation of this information into polypeptides. See Also

Polypeptides are proteins and enzymes that are
composed of amino acids that are polymerized ▶ Cell Wall
through peptide bonds. Enzymes catalyze most ▶ Exopolymers
metabolic reactions in the cell. Glycogen, ▶ Nucleic Acids
chrysolaminaran, and starch are examples of ▶ Peptidoglycan
branched and linear polyglucoses that serve as ▶ Polymer
carbon and energy storages in a variety of organ- ▶ Polynucleotide
isms. Polyhydroxyalkanoates (PHA), such as ▶ Polypeptide
polyhydroxybutyrate (PHB), also serve as ▶ Polysaccharide
reserve compounds in many bacteria. These com- ▶ Protein
pounds have found application as biodegradable ▶ Refractory Organic Polymer
plastics. Cyanophycin is a polymer also known as
multi-L-arginyl-poly-[L-aspartic acid]. It serves
as nitrogen storage, and although it was first
discovered in cyanobacteria (therefore the Decho AW (1990) Microbial exopolymer secretions in
name), it also occurs in other bacteria. This poly- ocean environments-their role(s) in food webs and
mer has found application as a pharmaceutical. marine processes. Oceanogr Mar Biol 28:73–153
Chitin is a linear homopolymer of N-acetyl- Joshi CP, Mansfield SD (2007) The cellulose paradox-
simple molecule, complex biosynthesis. Curr Opin
glucosamine. Chitin and cellulose (a glucose Plant Biol 10:220–226
polymer) share a similar structure. Chitin is a Latgé JP (2007) The cell wall: a carbohydrate armour for
component of the cell walls of arthropods and the fungal cell. Mol Microbiol 66:279–290
most fungi and is also synthesized by some dia- Passow U (2002) Transparent exopolymer particles (TEP)
in aquatic environments. Prog Oceanogr 55:287–333
toms, but is not known in bacteria or archaea. Rinaudo M (2006) Chitin and chitosan: properties and
Cellulose is a typical plant ▶ cell wall component applications. Prog Polym Sci 31:603–632
but is also synthesized by some bacteria, notably Verlinden RAJ, Hill DJ, Kenward MA, Williams CD,
by cyanobacteria. Peptidoglycan (murein) is a Radecka I (2007) Bacterial synthesis of biodegradable
polyhydroxyalkanoates. J Appl Microbiol 102:
biopolymer of N-acetyl-glucosamine and 1437–1449
N-acetyl-muramic acid, which serves as the
main cell wall component in bacteria and some
archaea (the latter possess a slightly different
polymer known as pseudopeptidoglycan). Bioprecipitation
Polyphosphate is an inorganic polymer of ortho-
phosphate and occurs in all three domains of life. Karim Benzerara
In bacteria, polyphosphates may serve as storage Institut de Minéralogie et de Physique des
of phosphorus and/or energy. Silicate is a mostly Milieux Condensés, UMR 7590, CNRS,
inorganic polymer of silicium although it is Université Pierre et Marie Curie & Institut de
normally associated with organic matter. Silicate Physique du Globe de Paris, Paris, France
is the main component of the frustules of diatoms.
Probably the most diverse and complex Synonyms
biopolymers are exopolymers. These are com-
plex polymeric carbohydrates with associated Biomineralization
glycolipids, proteins, and nucleic acids, as well
as a variety of uronic acids, pyruvate, sulfate, and
carboxylic groups. They may serve as the sheath Definition
of microorganisms or be extruded as mucilage in
the outer environment and have a variety of Bioprecipitation refers to the process of forma-
functions. tion of mineral phases (bioprecipitates or
Biosensor 311
biominerals) by organisms. Cells or associated the transducer, which delivers a signal propor-
extracellular polymers can serve as nucleation tional to the concentration of the substance (see
surfaces and/or alter the chemical composition Fig. 1). The main advantages of biosensors are
of fluids and raise the supersaturation of the solu- the selectivity and sensitivity of the analysis. In
tion with a mineral phase. Alternatively, addition, biosensors can supply real-time and B
bioprecipitation has been used in meteorology online information. Biological elements
to refer to the nucleation of ice by bacteria in employed can be enzymes, antibodies, DNA,
clouds resulting in snow or rainfall. whole cells, or tissues, for example. Depending
on the physical change observed, transducers
could be electrochemical, optical, piezoelectric,
acoustic, magnetic, or nanomechanical.
See Also
Overview
Biosensor is a general term for a wide range of

analytical devices that measure the presence or
Biosensor concentration of biological or chemical mole-
cules in complex samples by translating a biomo-
Laura M. Lechuga lecular interaction at the transducer surface into a
Nanobiosensors and Bioanalytical Applications quantifiable physical signal. When the
Group, Institut Català de Nanociència i biospecific interaction takes place, a physico-
Nanotecnologia (ICN2) CSIC and CIBER-BBN, chemical change is induced, as, for example, a
Barcelona, Spain redox, mass, resonant frequency, absorption, or
refractive index change. The aim of a biosensor is
to produce discrete or continuous signals that are
Keywords proportional to the concentration of a single ana-
lyte or a related group of analytes. The biological
Transducer; Bioreceptor layer; Optical biosen- element is capable of sensing the presence, activ-
sor; Electrochemical biosensor; Acoustic biosen- ity, or concentration of the substance in a solu-
sor; Plasmonic sensors; Label-free detection; Lab tion. Substances capable of being analyzed with
on a chip biosensors range from protein biomarkers,
toxins, antigens, drugs of abuse, pesticides, pep-
tides, proteins (IgG, albumin, insulin, etc.), hor-
Synonyms mones, vitamins, antibiotics, organic molecules
(glucose, lactose, cholesterol, urea, etc.), DNA
Biodetection system; Biological sensors; Lab on and DNA single mutations, RNA, microRNA,
a chip epigenetic marks, cancer cells, pathogenic micro-
organisms, and many others.
The first demonstration of a biosensor dates
Definition from 1962, when enzyme electrodes for glucose
sensing were developed by L. C. Clark (USA).
A biosensor is a compact hybrid device incorpo- Since then, an intensive effort has been focused
rating a biological receptor in direct contact with on biosensor research and development, a signif-
a transducer (the term “biosensor” derives from icant output of which was the commercial launch
this combination). The biological element specif- of the glucose biosensor in 1975. The glucose
ically recognizes a substance (chemical or bio- biosensor has become a landmark in the diagnos-
logical) in a sample, thereby inducing a change in tics field, thereby allowing millions of diabetics
312 Biosensor
Biosensor, Fig. 1 Scheme of a biosensor device
to test, in any place and at any time, their glucose measurable signal, and (ii) the stable and reliable
level in the blood. Substantial work on achieving biofunctionalization of the physical transducer
reliable biosensors began in the 1980s, and since with the appropriate, selective, biological recep-
then, many developments have been demon- tor (protein, DNA, cell, aptamer, etc.). The bio-
strated. Biosensing technology has been a very sensor research and development area is highly
active research field both in academic and in multidisciplinary because several aspects need to
industrial laboratories during the last decades. be considered to successfully achieve the device,
But, until now, only a limited number of biosen- such as the transduction modality used, fluidic
sors have achieved the stringent requirement for design (most of the biosensing analyses are
being commercialized. This is mainly due to the performed in liquid), surface biofunctiona-
poor stability of the biological receptor once it is lization, detection format (direct, indirect,
in contact with the transducer, as it must be stable sandwich-type, competitive binding), and data
under variable conditions of pH, temperature, processing and analysis.
humidity, and ionic strength.
The use of biosensor devices has clear advan-
tages in analytical chemistry. The main charac- Basic Methodology
teristics of biosensors are high sensitivity and
selectivity. In addition, biosensors can provide The possible combinations of different
rapid, direct, and cost-effective analyses, ease of bioreceptors and transducers result in a vast
operation, high accuracy, and wide detection range of biosensor types. Depending on the
range using a minimum volume of sample and bioreceptor, the transducer, or the type of detec-
reagents. This technology avoids the expensive, tion, biosensors can be classified into different
complex, and time-consuming procedures typi- types and subtypes. It is important to point out
cally employed in standard analytical evaluation. that the selection of the most suitable combina-
In order to obtain reliable biosensor devices, tion of bioreceptor/transducer/type of detection is
there are two crucial steps: (i) the design and dictated by the application itself, which depends
fabrication of a highly sensitive physical trans- on the type of substance to be detected and the
ducer, that is, a device capable of efficiently complexity of the sample which contains the
transforming the biomolecular reaction into a substance to be evaluated.
Biosensor 313
Types of Detection biosensing reaction involves redox changes.

Two main types of detection can be employed: They are the simplest and most developed bio-
“label” and “label-free.” In the “label-free” sensors. They can be amperometric (based on the
scheme, the original and unmodified analyte is measurement of the current resulting from the
detected in a direct manner. In the “label” electrochemical oxidation or reduction of B
approach, the analyte is tagged with a label electroactive species), potentiometric (based on
which acts as an indirect indicator of the presence the determination of the potential difference
of the molecule. In general, the “label” approach between an indicator and a reference electrode
results in a higher sensitivity of analysis, while (such as ISFET devices)), or conductometric
the “label-free” approach is direct, is rapid, and (based on a change in conductance (such as mea-
avoids the manipulation or degradation of the sured by interdigitated electrodes)). Piezoelectric
analyte. Each detection method has advantages biosensors are based on the piezoelectric effect:
and disadvantages and the method employed the frequency of a resonating crystal changes
should be selected according to the foreseen when molecules are adsorbed on top of the crys-
application. tal. Surface acoustic wave systems and quartz
crystal microbalances are the main examples of
Classification by Bioreceptor Layer Type piezoelectric devices.
Depending on the bioreceptor, biosensors can be In optical biosensors, the biorecognition event
divided into two main types: catalytic biosensors results in a change in optical properties such as a
and affinity biosensors. In catalytic biosensors, variation in the emission properties
the receptors are able to recognize (bio)chemical (luminescence), in the absorption coefficient, or
species and transform them into a product. This in the refractive index. This variation induces a
type of biosensor is mostly represented by enzy- change in some of the characteristics of light
matic biosensors, but whole cells or tissue recep- (wavelength, intensity, polarization, phase veloc-
tors also fall under this category. Affinity ity) which can be detected using techniques as
biosensors use the specific capabilities of an ana- emission, absorption, fluorescence, evanescent
lyte to bind to a biorecognition element. wave detection, or polarimetry. Many optical bio-
Immunosensors (based on specific interactions sensors have been developed because they fre-
between an antibody and an antigen) or DNA quently exhibit higher sensitivity than other
biosensors (based on the affinity between com- transducer types. One of the most successful
plementary oligonucleotides) are the main exam- label-free and commercially accepted optical
ples of affinity biosensors. Biomimetic receptors biosensors is the surface plasmon resonance
based on engineered molecules as aptamers or (SPR) sensor that is based on an evanescent
molecular imprinted polymers have emerged dur- wave interaction. Localized surface plasmon res-
ing the last years as an attractive alternative to onance (LSPR) based on metal nanostructures
biological receptors, although they have not has gained attention in the last years. Other opti-
achieved yet the same specificity than the natural cal biosensors make use of optical waveguides as
receptors and they are only suitable for some the basic element for light transmission, such as
specific applications. optical fibers or integrated optics-based wave-
guides, including interferometers (Mach-
Classification by Transducer Type Zehnder, Young, bimodal interferometers),
Depending on the transducer employed, biosen- microcavities (as microring resonators), and
sors can be divided into the following main types: grating- and photonic crystal-based structures.
electrochemical biosensors, optical biosensors, Other optical biosensors include those based on
piezoelectric biosensors, magnetic biosensors, porous silicon, silicon photonic wires, or resonant
and nanomechanical biosensors. Electrochemical bicells.
biosensors are based on changes in electrical In nanomechanical sensors, the specific
currents or potential across electrodes when the biorecognition on the cantilever surface results in
314 Biosensor
a nanomechanical response, which produces a can- research in the pharmaceutical industry, online
tilever bending of few nanometers and a shift in the screening of water quality for both household
resonant frequency. In this way, microcantilevers and industrial use, control of industrial processes,
translate the molecular reaction into a veterinary diagnostics, quality control in the food
nanomechanical motion, which is commonly and beverage industry, detection of harmful sub-
detected using optical or piezoresistive readout. stances, military and homeland security or envi-
Nanomechanical sensors achieve a high level of ronmental monitoring, and many others. The
sensitivity and are amenable for array multiplexing most significant applications are related to the
and integration in compact platforms. clinical diagnostics field, mainly due to the great
success of glucose biosensors since their inven-
Biofunctionalization tion. Indeed, glucose biosensors account for
Biofunctionalization is the process of about 85 % of the entire biosensor market. In
immobilizing the receptor biomolecules onto the astrobiology area, biosensors are useful tools
the transducer surface to provide the re- for detecting biomarkers or biosignatures of
quired biospecificity of the biosensor. The extant or extinct life which could be produced in
bio-immobilization process must maximize the biotic or abiotic processes (such as bacteria
sensitivity and must supply a reproducible layer strains, cells, extracellular materials, isolated
and must ensure optimal coverage density of the macromolecules or biomarkers, etc.).
receptor, proper receptor orientation for efficient
kinetics, minimal background and nonspecific
adsorption, and long-term stability. Amongst the Future Directions
various bio-immobilization approaches, the most
commonly used are physical adsorption, covalent The ultimate aim in biosensor technology is the
binding, and affinity-based (as avidin/biotin, achievement of a fully integrated, low-cost, por-
His-Tag systems, protein A/G for antibodies) table, and reliable lab-on-a-chip platform, able to
entrapment (physical or covalent) within host simultaneously detect and identify hundreds of
matrices (e.g., polymers, polymeric hydrogels, substances in real time with high sensitivity, even
and sol-gels). Usually, covalent binding to the at the single molecule level and connected
substrate is the preferred route for most through a wireless scheme to a central opera-
bioreceptors through the use of self-assembled tional station or hospital database. Therefore,
monolayers, but the most suitable technique the major future objectives in biosensor technol-
depends on the bioreceptor type and the sample ogy will be to achieve higher sensitivity, to
to be evaluated. reduce sample volume and reagents by miniatur-
Whatever the immobilization technique ization at the micro-/nanoscale, to increase the
employed, a previous chemical activation of the number of substances detected within the same
sensor surface is mandatory. Silicon-, silicon sample, to develop devices of high multiplexing
oxide-, and silicon nitride-based surfaces can be capabilities for analyzing an elevated number of
functionalized using the well-known silane substances in parallel, and to develop wearable
chemistry. Gold surfaces are normally biosensors and biosensors suitable for implanta-
functionalized using thiol chemistry. tion in the human body. Other important aspects
to be considered in the future are issues associ-
ated with the commercialization of biosensors for
Applications relevant applications, which still face significant
technological hurdles. Development cycles tend
The fields of application for biosensors are almost to be long and commercial success remains lim-
endless. Applications for these devices include ited. The main issues in biosensor technology,
clinical diagnostics (including personal health such as the stability and reliability of the
monitoring), basic research in life sciences, drug biorecognition layer, still need to be surpassed.
Biosignatures, Effect of Metamorphism 315
See Also The 2009–2014 World Outlook for Biosensors

(2009) ICON Group International, Inc.
Vollmer F, Yang L (2012) Label-free detection with high-
▶ Aptamer Q microcavities: a review of biosensing mechanisms
▶ Hybridization for integrated devices. Nanophotonics 1(3–4):267–291
▶ Molecular Recognition Wang J (2008) Electrochemical glucose biosensors. Chem B
▶ Surface Plasmon Resonance Rev 108:814–825
Wang P, Liu Q (eds) (2009) Cell-based biosensors: prin-
ciples and applications, 1st edn, Engineering in medi-
cine & biology. Artech House, New York
Alvarez M, Lechuga LM (2010) Microcantilever-based

platforms as biosensing tools. Analyst 135:827–836
Borisov SM, Wolfbeis OS (2008) Optical biosensors. Biosignature
Chem Rev 108:423–461
Comeaux R, Novotny P (2009) Biosensors: properties,
materials and applications, Biotechnology in agricul- ▶ Biogenicity
ture, industry and medicine series. Nova Science Pub- ▶ Biomarkers
lishers, New York ▶ Biomarkers, Morphological
Cooper MA (ed) (2009) Label-free biosensors: techniques
and applications. Cambridge University Press,
Cambridge
Estevez MC, Alvarez M, Lechuga LM (2012) Integrated
optical devices for lab-on-a-chip biosensing applica- Biosignatures, Effect of
tions. Laser Photonics Rev 6(4):463–487 Metamorphism
Fan X, White IM, Shopova SI, Zhu H, Suter JD, Sun
Y (2008) Sensitive optical biosensors for unlabeled
targets: a review. Anal Chim Acta 620:8–26 Sylvain Bernard
Goeders KM, Colton JS, Bottomley LA (2008) Microcan- Laboratoire de Minéralogie et de Cosmochimie
tilevers: sensing chemical interactions via mechanical du Muséum (LMCM), Paris, France
motion. Chem Rev 108:522–542
Homola J (ed) (2006) Surface plasmon resonance based
sensors, 1st edn, Springer series on chemical sensors
and biosensors. Springer, Berlin Keywords
Homola J (2008) Surface Plasmon resonance sensors for
detection of chemical and biological species. Chem
Rev 108:462–493
Burial; Chemistry; Degradation; Diagenesis;
Länge K, Rapp BE, Rapp M (2008) Surface acoustic wave Fossilization; Isotopes; Metamorphism; Micros-
biosensors: a review. Anal Bioanal Chem copy; Organic molecules; Preservation; Spec-
391:1509–1519 troscopy; Structure
Ligler F (ed) (2008) Optical biosensors: today and tomor-
row, 1st edn. Elsevier Science, Amsterdam
Marks RS, Lowe CR, Cullen DC, Weetall HH, Karube
I (eds) (2007) Handbook of biosensors and biochips. Synonyms
Wiley-Interscience, Chichester
Privett BJ, Shin JH, Schoenfisch MH (2008) Electrochem-
ical sensors. Anal Chem 80:4499–4517 Fossilization processes
Rasooly A, Herold KE (eds) (2009) Biosensors and
biodetection: 1 (methods in molecular biology):
methods and protocols, volume 1: optical-based detec- Definition
tors and volume 2: electrochemical and mechanical
detectors, lateral flow and ligands for biosensors.
Humana Press Springer Protocols, Totowa Biosignatures are molecular, mineral, or isotopic
Sepúlveda B, Angelomé PC, Lechuga LM, Liz-Marzán patterns that can be unambiguously interpreted as
LM (2009) LSPR-based nanobiosensors. Nano Today evidence of extant or extinct life (Slater 2009).
4:244–251
Steinem C, Janshoff A (eds) (2007) Piezoelectric sensors, However, fossilized biosignatures are not immu-
1st edn, Springer series on chemical sensors and bio- table but prone to degradation during diagenesis
sensors. Springer, Berlin and ▶ metamorphism and may become difficult
316 Biosignatures, Effect of Metamorphism
to distinguish from signatures of abiogenic com- In the long term, physical biosignatures have a
pounds (e.g., Garcı́a-Ruiz et al. 2003; Pasteris greater chance of preservation than complex
and Wopenka 2003; McCollom and Seewald organic markers. Metamorphic degradation of
2006). Nevertheless, from the multiscale charac- ▶ organic molecules is associated with an aroma-
terization of organic-rich ▶ metamorphic rocks ticity increase, mainly by the release of
using advanced synchrotron-based micro ▶ spec- heteroelement-containing functional groups as
troscopy techniques, Bernard et al. (2007, 2010) CO2, N2, and H2S and by the elimination of ali-
have shown that high-grade metamorphism phatic groups as CH4. Organic molecules which
might not totally erase structural and chemical have undergone restructuring during early diagen-
bio-features, at least at the sub-micrometer scale. esis to become resistant cross-linked aliphatic or
aromatic macromolecules (a.k.a. geopolymers)
and those which have been closely associated to
Overview minerals (such as carbonates, sulfates, and clays)
will have a greater chance of long-term preserva-
Understanding the physical and chemical evolution. The isotopic composition of organic com-
tion of life relies on the ability to decipher pre- pounds is relatively stable, to the extent that basic
served evidence in the rock record, a.k.a. molecular skeletons are preserved. Otherwise, the
fossilized biosignatures (Seckbach and Walsh effect of thermal metamorphism on organic matter
2008; Slater 2009). However, the combination is to form isotopically lighter volatile species and
of biological, chemical, and physical factors dur- isotopically heavier residual refractory solids. In
ing taphonomic processes and deep burial inevi- conclusion, although biosignatures might be par-
tably alter the chemical, structural, and isotopic tially preserved during metamorphism, a great deal
fidelity of such fossilized biosignatures of caution must be implemented and multiple lines
(Schiffbauer et al. 2007). In addition, abiogenic of evidence (morphological, ultrastructural, as well
processes may form disordered carbon- as geochemical) must be sought during their anal-
dominated precipitates displaying morphologies ysis and interpretation.
and signatures quite similar to biogenic objects
(Garcı́a-Ruiz et al. 2003; Pasteris and Wopenka
2003; McCollom and Seewald 2006). Therefore, See Also
unambiguously identifying biosignatures within
metamorphic rocks has been generally believed ▶ Abiotic
to be extremely difficult, if not impossible, except ▶ Absorption Spectroscopy
in very rare cases such as the exceptionally well- ▶ Biomarkers
preserved Burgess Shale ▶ fossils which experi- ▶ Biomarkers, Morphological
enced greenschist facies metamorphism (Powell ▶ Biomarkers, Spectral
2003). Recently, the analytical improvements of ▶ Biopolymer
microspectroscopy techniques in general and ▶ Carbon Cycle, Biological
synchrotron-based techniques in particular (see ▶ Carbon Isotopes as a Geochemical Tracer
Templeton and Knowles 2009 for a review) ▶ Complex Organic Molecules
have allowed the chemical and structural charac- ▶ Fischer-Tropsch Effects on Isotopic
terization of metamorphic organic matter Fractionation
down to the nanometer scale. Altogether, these ▶ Fischer-Tropsch-Type Reaction
studies have shown that even high-grade meta- ▶ Fossil
morphic rocks may retain morphologically and ▶ Fossilization, Process of
geochemically recognizable traces of life, ▶ Infrared Spectroscopy
depending notably on the original biochemical ▶ Isotope
nature of the ▶ biopolymer (Bernard et al. 2007, ▶ Isotope Biosignatures
2010). ▶ Isotopic Exchange Reaction
Biosphere 317

▶ Isotopic Ratio Biosphere
▶ Metamorphic Rock
▶ Metamorphism David C. Fernández-Remolar
▶ Metasediments Centro de Astrobiologı́a (INTA-CSIC), INTA, B
▶ Microfossils Torrejón de Ardoz, Spain
▶ Microfossils, Analytical Techniques
▶ Organic Molecule Keywords
▶ Origin of Life
▶ Raman Spectroscopy Biocenosis; Biogeochemistry; Ecosystem;
▶ Spectroscopy Homeostasis
References and Further Reading Synonyms

Bernard S, Benzerara K, Beyssac O, Menguy N, Guyot F, Ecosphere
Brown GE Jr, Goffe B (2007) Exceptional preserva-
tion of plant fossils in high-pressure metamorphic
rocks. Earth Planet Sci Lett 262(1–2):257–272
Bernard S, Benzerara K, Beyssac O, Brown GE Jr Definition
(2010) Multiscale characterization of pyritized plant
tissues in blueschist facies metamorphic rocks.
Biosphere is a term initially created by the prom-
Geochim Cosmochim Acta 74:5054–5068
Garcı́a-Ruiz J, Hyde S, Carnerup A, Christy A, Van inent geologist Edward Suess to define the upper
Kranendonk M, Welham N (2003) Self-assembled part of the Earth’s lithosphere that is intricately
silica-carbonate structures and detection of ancient associated to life (Smil 2003). Later on, the Rus-
microfossils. Science 302:1194–1197
sian biologist Vladimir Vernadsky realized the
McCollom TM, Seewald JS (2006) Carbon isotope com-
position of organic compounds produced by abiotic real dimension of the impact of life on Earth
synthesis under hydrothermal conditions. Earth Planet and expanded the biosphere to the global geo-
Sci Lett 243(1–2):74–84 chemical cycling of matter within the terrestrial
Pasteris JD, Wopenka B (2003) Necessary, but not suffi-
crust (Vernadsky 1998). According to this last
cient: Raman identification of disordered carbon as a
signature of ancient life. Astrobiology 3(4):727–738 conception, biosphere can be understood as a
Powell W (2003) Greenschist-facies metamorphism of the complex, natural entity resulting, on a global
Burgess Shale and its implications for models of fossil scale, from the interactions between all life
formation and preservation. Can J Earth Sci 40(1):13–25
forms and the geosphere, ▶ hydrosphere, and
Schiffbauer JD, Yin LM, Bodnar RJ, Kaufman AJ, Meng
FW, Hu J, Shen B, Yuan XL, Bao HM, Xiao SH atmosphere of a planetary body (Smil 2003).
(2007) Ultrastructural and geochemical characteriza- Based on this concept, the British scientist
tion of archean-paleoproterozoic graphite particles: James Lovelock envisaged the biosphere as an
implications for recognizing traces of life in highly
organism of planetary scale, Gaia, which rules the
metamorphosed rocks. Astrobiology 7(4):684–704
Seckbach J, Walsh M (2008) From fossils to main physical and chemical parameters of the
astrobiology – records of life on Earth and the search planet through a series of intricate feedback
for extraterrestrial biosignatures. In: Cellular origin, mechanisms (Lovelock 2000).
life in extreme habitats and astrobiology. Springer
science series, 12. p 548
Slater GF (2009) Biosignatures: interpreting evidence of the
origins and diversity of life. Geosci Can 36(4):170–178 Overview
Templeton A, Knowles E (2009) Microbial transforma-
tions of minerals and metals: recent advances in
Under the formal ecological terminology, the
geomicrobiology derived from synchrotron-based
x-ray spectroscopy and x-ray microscopy. Annu Rev biosphere can be defined as a global biological
Earth Planet Sci 37:367–391 system that arises from the integration of all
318 Biosphere
biocenosis (Lévêque 2003), ranging from those specialized colonists and ends in a highly diverse
sustained by primary photosynthesizers to that ecosystem sustained by very specialized organ-
uniquely maintained by chemolithotrophic pro- isms adapted to a wide variety of niches. To the
ducers as in deep crustal regions. In this sense, the same extent, but at greater scale, and coupled to
ecosphere would include the biosphere as the the evolution of the biosphere, oceanic and atmo-
biotic component of the planetary ▶ ecosystem, spheric oxidation was promoted by bacterial com-
in the same manner that biocenosis accounts for munities (cyanobacteria) releasing oxygen into the
ecosystem (Huggett 1999). As it is difficult to ocean and the atmosphere, where it was used by
separate biology from the environment with more complex microorganisms that were able to
which it interacts, the biosphere is currently iden- use oxygen for respiration, which in turn
tified by its interactions with the different plane- unleashed the emergence of the multicellular
tary envelopes such as the atmosphere, the organisms. As a consequence, microbial-mediated
hydrosphere, or the crust. As a consequence of oxygenation in combination with global geologi-
the continuous exchange of matter and energy cal events (Knoll 2003) triggered diversity and
between living forms and the planet, such inter- environmental change in the form of positive feed-
actions are materialized in the form of complex back on Early Earth that affected not only the
mechanisms that induce the control of the phys- ocean ▶ habitats but also led to the emergence of
ical and chemical variables of the environment on habitability on the continental areas of the Earth.
regional and global scales. One paradigmatic
example of this is the temperature and pH control
in the oceans (Orr et al. 2005; Knoll et al. 1996) See Also
through the uptake and sequestration of CO2,
which is transformed into biomass and carbon- ▶ Algae
ates, which are later buried. The same kind of ▶ Archaea
global feedbacks have been observed in other ▶ Autotroph
compounds bearing nitrogen and sulfur, which ▶ Biogeochemical Cycles
can induce physical and chemical changes in the ▶ Biotope
Earth’s atmosphere after being included in the ▶ Carbon Cycle, Biological
photochemical pathways of the troposphere ▶ Earth’s Atmosphere, Origin and Evolution of
(Seinfeld and Pandys 2006). Such processes pro- ▶ Ecosystem
voke thermal changes on the Earth’s surface by ▶ Gaia Hypothesis
increasing the planetary albedo as SO2 currently ▶ Greenhouse Effect
does or reinforcing the ▶ greenhouse effect as ▶ Habitat
occurs with some N-bearing gases. Moreover, the ▶ Heterotroph
manner in which the structure, composition, and ▶ Homeostasis
substrate of extensive woodlands control the water ▶ Hydrosphere
and thermal balance, which is essential for hydro- ▶ Nitrogen Cycle, Biological
logical cycles at regional and global scales, has ▶ Ocean, Chemical Evolution of
been repeatedly described (Rodrı́guez-Iturbe and ▶ Oxygenation of the Earth’s Atmosphere
Porporato 2004). Interestingly, the great oxygena- ▶ Sulfur Cycle
tion event driven by microbial activity during the ▶ Symbiosis
Late Archean-Early Proterozoic can be consid-
ered, on a greater scale, as a similar process of
environmental control leading to the emergence of
▶ homeostasis as observed in the ecological suc- Huggett RJ (1999) Ecosphere, biosphere or Gaia? What to
cession of areas, which are initially devoid of call the global ecosystem. Glob Ecol Biogeogr
living forms. This process starts with few low 8(6):425–431
Biostack 319
Knoll AH (2003) Life on a young planet: the first three oriented horizontally, and EPS (extracellular
billion years of evolution on earth. Princeton Univer- polymeric substances, if present) change their
sity Press, Princeton
Knoll AH, Bambach RK, Canfield DE, Grotzinger JP chemical structure to a more erosion-resistant
(1996) Comparative earth history and late Permian phase. Biostabilization by epibenthic and
mass extinction. Science 273:452–457 endobenthic microbial mats is expressed by B
Lévêque C (2003) From ecosystem to biosphere. Science
Publishers, Enfield
Lovelock J (2000) Gaia, a new look at life on earth, Y ¼ ru2 =ðrs rf ÞgDn
4th edn. Oxford University Press, Oxford
Orr JC, Fabry VJ, Aumont O, Bopp L, Doney SC, Feely where u is the shear velocity, rf is the density of
RA, Gnanadesikan A, Gruber N, Ishida A, Joos F, Key
fluid, rs is the density of sediment, g is the gravity
RM, Lindsay K, Maier-Reimer E, Matear R,
Monfray P, Mouchet A, Najjar RG, Plattner G-K, constant, D is the actual grain diameter under the
Rodgers KB, Sabine CL, Sarmiento JL, Schlitzer R, influence of biostabilization, and n is the expo-
Slater RD, Totterdell IJ, Weirig M-F, Yamanaka Y, nent to which D is raised for the data to comply to
Yool A (2005) Anthropogenic ocean acidification over
the Shield’s relationship. Type 2 biostabilization
the twenty-first century and its impact on calcifying
organisms. Nature 437:681–686 is the flexible deformation of otherwise brittle
Rodrı́guez-Iturbe I, Porporato A (2004) Ecohydrology of sediment composed of loose sand grains. Type
water-controlled ecosystems. Cambridge University 3 biostabilization is sealing of the sedimentary
Press, Cambridge, UK
surface so that exchange of gases between sedi-
Seinfeld JH, Pandys SN (2006) Atmospheric chemistry
and physics, 2nd edn. Wiley, New Jersey ment and atmosphere or water is prohibited.
Smil V (2003) The earth’s biosphere: evolution, dynamics
and change. The MIT Press, Cambridge, MA
Vernadsky VI (1998) The biosphere (annotated and
revised by Mark A. S. McMenamin). Copernicus,
See Also
Springer-Verlag, New York
▶ Biofilm
▶ Microbial Mats
▶ Microbially Induced Sedimentary Structures
▶ MISS
Biostabilization
Nora Noffke
Department of Ocean, Earth & Atmospheric Biostack
Sciences, Old Dominion University, Norfolk,
VA, USA Gerda Horneck
DLR German Aerospace Center, Institute
of Aerospace Medicine, Radiation Biology,
Synonyms Köln, Germany
Microbial sediment fixation

Keywords
Definition Biological radiation effects; Cosmic rays; Heavy

ion hits; HZE particles; Nuclear track detectors
Biostabilization is fixation of sedimentary grains
by ▶ biofilms and ▶ microbial mats. Three types
of biostabilizations are distinguished. Type Synonyms
1 biostabilization is the response by benthic
microbiota to erosion. Bacterial filaments are Biobloc
320 Biostack
Definition radiation interact with biological systems, the

Biostack concept was developed to precisely
The Biostack concept allows studying the biolog- localize the trajectory of an HZE particle relative
ical effects of single heavy ions of cosmic rays. to the biological object and to correlate the phys-
Each Biostack device is composed of a stack of ical data of the particle relative to the observed
visual track detectors (e.g., nuclear emulsions, biological effects along its path (Fig. 1). A large
cellulose nitrate, and polycarbonate) with layers variety of biological specimens, such as viruses,
of biological objects in resting state sandwiched bacterial ▶ spores, plant seeds and animal eggs,
between them. After exposure to the outer space, and cysts, allowed the evaluation of radiation
the track detectors are developed, the trajectory effects at different levels of biological organiza-
of each heavy ion is localized relative to the site tion and at different radiation sensitivities. The
of the biological object, and the physical data of following biological effects were ascribed to the
each particle are correlated with the observed passage of a single HZE particle:
biological effects along its path.
• In seeds of Arabidopsis thaliana or Nicotiana
tabacum, hit by an HZE particle in their shoot
History meristem, development was significantly dis-
turbed, as demonstrated by up to 90 % loss of
The Biostack program has been developed in the germination (early lethality) or embryo lethal-
late 1960s under the auspices of the Council of ity. Seedling abnormalities, such as hypertro-
Europe within the Working Group on Space Bio- phy or deformation of cotyledons, hypocotyl,
physics and has been performed in international or root, or chlorophyll deficiency, occurred
cooperation by scientists from Europe, Russia, with high frequency as a consequence of a
and the USA. Biostack experiments were flown passage of a single HZE particle close to the
on the NASA missions Apollo 16 and 17 (1972), shoot or root meristem. In Lactuca sativa
Apollo-Soyuz Test Project (1975), and the seeds, hit by an HZE particle, multiple chro-
▶ Long Duration Exposure Facility mosomal aberrations developed at high fre-
(1984–1990), on ESA/NASA missions Spacelab quencies. Zea mays seeds, flown on ASTP,
1 (1983) and Spacelab IML-2 (1994), on the developed large yellow strips in all leaves as
German/NASA mission Spacelab D2 (1993), on somatic mutation.
the ESA mission ▶ EURECA (1992–1993), and • In 90 % of the mosaic eggs of the brine shrimp
on ESA/Russian missions of the Biocosmos and Artemia salina hit by an HZE particle, hatch-
▶ Foton series (1992–1994). The Biostack pro- ing, characterized by release of a free-
gram was managed by Horst B€ucker and G€unther swimming nauplius, was inhibited. Anomalies
Reitz, both from the German Aerospace Center of the body or extremities appeared approxi-
(DLR), Germany. mately ten times more frequently than in the
ground controls. Likewise, hatching of the
Indian stick insect Carausius morosus from
Overview eggs, hit by a cosmic HZE particle, was sig-
nificantly reduced. Malformations were
With the beginning of human space flight, health increased in individuals having developed
effects from exposure to cosmic rays became a from eggs hit. They were characterized by a
concern, and radiation protection guidance has curved abdomen, fused segments, or
been required. The heavy ions of cosmic radia- shortened legs.
tion, the so-called ▶ HZE particles, i.e., particles • Spores of the bacterium Bacillus subtilis were
of high (H) atomic number (Z) and high energy inactivated well beyond 1 mm from the HZE
(E), are of special concern. To understand the particle trajectory, whose distance would
ways by which single particles of cosmic roughly correspond to the dimensions of a
Biostack 321
Biostack, Fig. 1 The Biostack concept to localize bio- Apollo 16 and 17; ASTP; Spacelab 1, D2, IML-1, and
logical effects produced by single HZE particles of cosmic IML-2; LDEF; Cosmos 1887 and 2004; and EURECA
radiation; Biostack experiments were flown onboard of (Photo credit DLR)
▶ spore, thereby pointing to a long-ranging ▶ Panspermia

effect of HZE particles. ▶ Radiation Biology
▶ Space Environment
Quantifying various genetic and developmen- ▶ Spore
tal disturbances caused by individual HZE parti-
cles of cosmic radiation, the Biostack data are of
relevance for assessing the radiation risks for References and Further Reading
astronauts, especially during exploratory mis-
sions, as well as for estimating the likelihood of B€
ucker H (1975) BIOSTACK – a study of biological
effects of HZE galactic cosmic radiation. In: Biomed-
▶ lithopanspermia, i.e., the interplanetary trans- ical results of Apollo, NASA SP-368. NASA, Wash-
fer of microorganisms within rocks. ington, DC, pp 343–354
B€
ucker H, Horneck G, Facius R, Reitz G, Schäfer M,
Schott JU, Beaujean R, Enge W, Schopper E,
Heinrich W, Beer J, Wiegel B, Pfohl R, Francois H,
See Also Portal G, Bonting SL, Graul EH, R€ uther W, Kranz AR,
Bork U, Koller-Lambert K, Kirchheim B, Starke ME,
▶ Apollo Mission Planel H, Delpoux M (1984) Radiobiological
▶ Cosmic Rays in the Heliosphere advanced Biostack experiment. Science 22:222–224
B€
ucker H, Facius R, Horneck G, Reitz G, Graul EH,
▶ EURECA Berger H, Höffken H, R€ uther W, Heinrich W,
▶ Foton Capsule, Spacecraft Beaujean R, Enge W (1986) Embryogenesis and
▶ HZE Particle organogenesis of Carausius morosus under spaceflight
▶ Ionizing Radiation, Biological Effects conditions. Adv Space Res 12(6):115–124
Facius R, Reitz G, Schäfer M (1994) Inactivation of indi-
▶ Linear Energy Transfer vidual Bacillus subtilis spores in dependence on their
▶ Lithopanspermia distance to single cosmic heavy ions. Adv Space Res
▶ Long Duration Exposure Facility 14(10):1027–1038
322 Biosynthesis
Horneck G (1992) Radiobiological experiments in space: specific assemblage of organisms. Biotope is

a review. Nucl Tracks Radiat Meas 20:185–205 almost synonymous with the term habitat, but
Horneck G (1994) HZE particle effects in space. Acta
Astronaut 32:749–755 while the subject of a habitat is a species or a
Horneck G, Baumstark-Khan C, Facius R (2006) Radia- population, the subject of a biotope is a biological
tion biology. In: Clément G, Slenzka K (eds) Funda- community.
mentals of space biology. Kluwer/Springer,
Heidelberg/Berlin/Dordrecht, pp 292–335
Swenberg CE, Horneck G, Stassinopoulos EG (eds)
(1993) Biological effects and physics of solar See Also
and galactic cosmic radiation, Part A, vol 243A
and B, NATO ASI series A: life sciences. Plenum, ▶ Colonization, Biological
New York
Biosynthesis
Bipolar Flow
▶ Anabolism
Steven W. Stahler
Biotic Crisis
▶ Mass Extinctions Keywords
Molecular flow; Star formation
Biotic Isotope Fractionation

Definition
▶ Isotope Biosignatures
Bipolar flows are a phenomenon associated with
young stars still embedded in their parent
▶ molecular clouds. There are two kinds of
Biotope flows – jets and molecular outflows. The first
represent high-speed stellar wind gas, ejected
Ricardo Amils into two narrow streams. The second are broader
Departamento de Biologia Molecular, cones of more slowly moving cloud gas. Each
Universidad Autónoma de Madrid, Madrid, molecular outflow is driven by a jet running
Spain down its central axis. What is seen most promi-
nently in the jets are shock waves created when
the wind runs into either ambient cloud gas or
Synonyms slower gas ejected earlier. Neither the launching
of the wind nor its collimation into jets is well
Environment; Habitat understood.
Definition Overview
A biotope is an area of homogeneous environ- It has long been known that massive stars emit
mental conditions providing living space for a strong winds that drive back nearby gas. Since
Bipolar Flow 323
these objects are all relatively young, there is represent cloud material being pushed outward
often such a gaseous environment, the remnant by the jet. This relatively cold gas is observed
of the cloud that formed the star itself. In the through the radio emission of molecules in the
1980s, astronomers were surprised to discover cloud. The dominant species, molecular hydro-
that even young stars of solar mass and below gen, is a weak emitter at these low temperatures, B
drive vigorous outflows. Each outflow appears as so astronomers generally utilize CO, which has a
two opposite streams (hence “bipolar”) emanat- strong spectral line at a wavelength of 2.6 mm.
ing from the central object. In the youngest exam- Smaller amounts of much warmer gas have been
ples, the star is so embedded in ambient dusty gas detected using other tracer molecules, such as
that it is optically invisible. Even in such cases, ammonia.
however, bipolar outflows are present. The flows Since stellar jets glow mainly at optical wave-
probably play a key role in clearing away lengths, they are easily obscured by dust in the
remaining material in star-forming molecular molecular clouds that lie near the driving stars.
clouds. Despite extensive study, the physical To see a jet, the star must be near the surface of
mechanisms underlying both their launching the cloud. Sometimes, the cloud has been par-
and collimation remain mysterious. tially ablated by ultraviolet radiation from a
Flows generated by young stars have two nearby massive star, revealing the young,
components – jets and molecular outflows. Both low-mass star and its jet. The millimeter radio
have bipolar morphologies. Otherwise, their emission used to detect molecular outflows, how-
properties are very different, undoubtedly ever, easily penetrates interstellar dust. Thus,
reflecting their distinct origins. Stellar jets consist these broader flows are relatively easy to find,
of gas emitted at high speed into a narrow open- and hundreds of examples exist. In contrast,
ing angle. A typical jet speed is 300 km/s, which there are still only a few dozen clear detections
is also the magnitude of the escape velocity from of stellar jets. The red-shifted arm of the jet,
the stellar surface. The gas has a temperature of which is penetrating back into the molecular
10,000 K. What we observe most prominently are cloud away from the observer, is often consider-
shock waves created when this high-speed gas ably fainter but has been detected in most cases.
rams into other material. These shock waves, The high-speed material in stellar jets must
which radiate in a variety of spectral emission represent the actual wind being emitted by the
lines, are called Herbig-Haro objects. Discovered star or the inner portion of its circumstellar disk.
in the 1950s, the shocks appeared as isolated, The rough match of the wind speed and the stellar
glowing patches. Thirty years later, astronomers escape velocity occurs in other examples of
detected narrow, faint strands linking the bright winds from stars over a broad range of masses
knots and leading back to a young, embedded and ages. How the wind is launched in this par-
star. Each arm of a stellar jet has a length of ticular case is not clear. The wind from our Sun is
about 0.1 pc. Individual Herbig-Haro objects pushed outward by the hot gas at the base of the
can be detected much farther out, up to several solar corona. The mass transport rate in a young
parsecs from the driving source. stellar jet is higher by 6 orders of magnitude, and
Molecular outflows are broader swathes of thermal pressure is inadequate. It is generally
much slower and cooler gas. They exhibit the agreed that the wind is tapping rotational energy.
same bipolar morphology and axis orientation The star, which is indeed rapidly rotating, also
as jets. Indeed, it is now believed that every contains a strong magnetic field. The wind could
molecular outflow has a jet running down its be flung out along the magnetic field lines, a
center. Gas in the broader outflows moves at process which also occurs in our Sun. If this is
speeds of typically 10 km/s, and most of its the basic process, then the jet is also carrying off
mass has a temperature of about 10 K. Since angular momentum, causing the star to slow its
this temperature also characterizes molecular rotation. The outflow could also facilitate accre-
clouds, the broader outflows are thought to tion of inner disk material onto the star.
324 Bipolar Flow
The bright knots (Herbig-Haro objects) An alternative notion is that the expanding jet
observed along jets mark locations where high- creates shocks in the surrounding molecular gas.
speed gas encounters material moving at lower These so-called reflecting shocks effectively
velocity. But what exactly is this target material? bend the flow back toward the axis. The converg-
In a few cases, it is clear that wind gas is ing flow then creates another pair of shocks and
impacting the ambient molecular cloud. There diverges outward once more. Alternating pairs of
are some well-documented examples, however, reflecting shocks are commonly seen in super-
of the knots being spaced symmetrically around sonic flows on Earth, such as the exhaust of a
the central star. No external cloud could possess rocket.
such a high degree of symmetry. Hence, most It is clear, in any case, that the jets interact
shocks must arise in a different way. strongly with molecular gas. The outflows are
An important clue comes from observing the observed to be clumpier than normal cloud gas.
space motion of Herbig-Haro objects; images Their speed also increases closer to the jet axis.
taken a few decades apart show a marked shift, Terrestrial jets create not only a sequence of shocks
indicating speeds of several hundred kilometers but also a collar of turbulent gas that widens until
per second. This speed is similar to that detected the central jet terminates. An analogous process
in the main body of the jet, which we have said could be occurring near young stars. Here, turbu-
represents stellar wind material. On the other lence might be induced by jet material spewing out
hand, the spectral emission lines from the knots transversely from each shock. Indeed, careful
indicate that the shocked material is largely neu- observation of Herbig-Haro shocks does show
tral so that gas enters the shock front at speeds of material ejected in this manner, with the hot spray
only about 30 km/s. of ionized gas creating a cocoon surrounding the
The explanation is that incoming material is shock. This spray could also be responsible for
hitting not a static environment but one already stirring up the surrounding cloud gas and dragging
in motion. That is, the shocks arise when rela- it forward as a molecular outflow.
tively fast wind gas encounters slower gas that However, interstellar jets can extend far
was emitted earlier. Thus, the pattern of knots beyond the turbulent, clumpy outflow. Astrono-
effectively constitutes a record of temporal wind mers have detected a number of stellar jets
variations, and the symmetrical patterns extending over a parsec from the central star,
observed are neatly explained. The inferred fluc- with little or no molecular gas. These long chains
tuations in wind speed are of order 10 %, with of Herbig-Haro objects are gently curved,
associated periods ranging from decades to reflecting the rotation or perhaps precession of
millennia. the underlying jet source. The period of rotation
The most mysterious aspect of stellar jets is is of order 10,000 years. Despite the fact that they
their remarkable degree of collimation. Most jets are traveling through nearly empty space, the
have an aspect ratio (length to width) of about shocks are slowing down. It is possible that
100 to 1. One popular idea is that the rotating momentum is being continually deposited in the
wind material twists up in its internal magnetic lateral spray of gas. Indeed, small clumps of gas
field. A twisted field exerts an inward force along are sometimes observed off to the side of the
the central axis. Perhaps this force keeps the jet main jet flow.
from diverging. On the other hand, twisted mag- While much of the underlying physics is still
netic fields created in the laboratory are unstable murky, the phenomenon of bipolar flows has
to reconnection. So are the twisted flux loops that clear importance in the process of star formation.
protrude from the solar surface; their First, solar-type stars form inside relatively small
reconnection gives rise to X-ray flares. molecular clouds called dense cores. Since only
Researchers are using numerical simulations to about a third of the core mass winds up in the star,
see if the more vigorous outflows along interstel- the rest must somehow be dispersed. Winds are
lar magnetized jets are more stable. the obvious candidate. On a larger scale, stars are
Birthline 325
born not as isolated objects but in groups. Ini- Definition

tially, these clusters are embedded within large
molecular clouds. Many such primitive groups After they have acquired most of their mass, but
have been detected by infrared telescopes. before they begin to fuse hydrogen, stars slowly
When the group first appears in optical light, the contract. In the HR diagram, these ▶ pre-main- B
stars are a few million years old, and there is still sequence stars descend along paths whose loca-
a substantial amount of ambient molecular gas. tion depends on the stellar mass. All paths start at
Again, these so-called T associations are com- the birthline, a curve lying above and roughly
monly observed. On the other hand, we never parallel to the main sequence. When the stars
see such groups with ages as high as 10 million within young clusters are placed in the HR dia-
years. The reason must be that the stars them- gram, they lie between the birthline and the main
selves have driven off the molecular cloud and sequence. The two curves intersect at a mass of
then dispersed into the field population. The obvi- about ten solar masses. Stars of greater mass
ous driving mechanism is ▶ stellar winds. In this ignite hydrogen while they are still ▶ protostars.
view, molecular outflows represent the dispersal If a young cluster contains such objects, they lie
of cloud gas, the key step in terminating star on the main sequence.
formation on both large and small scales.
Overview
See Also
Astronomers view the evolution of stars using
▶ Pre-main-sequence Star the Hertzsprung-Russell (HR) diagram. Within
▶ Protostars this plot of stellar luminosity versus surface
▶ Stellar Winds temperature, the point representing a star
moves about as the object evolves. Mature
stars that are fusing hydrogen lie along the
so-called main sequence. Pre-main-sequence
Arce HG, Shepherd D, Gueth F, Lee C-F, Bachiller R, stars are situated higher in the diagram and
Rosen A, Beuther H (2007) In: Reipurth B, Jewitt D, gradually descend to the main sequence. The
Keil K (eds) Protostars and planets V. University of actual path of descent, known as the pre-main-
Arizona Press, Tucson, p 245 sequence track, depends on the stellar mass. All
Bachiller R (1996) Annu Rev Astron Astrophys 34:121
Reipurth B, Bally J (2001) Annu Rev Astron Astrophys pre-main-sequence tracks begin on another
39:403 curve known as the birthline.
Stahler SW, Palla F (2004) The formation of stars. Wiley- This description of the birthline is essentially
VCH, Weinheim, chapter 13 theoretical. Observationally, young ▶ stellar
clusters are studied by placing their members in
the HR diagram (also called the color-magnitude
diagram in this context). The birthline is the upper
Birthline envelope of the stellar distribution. That is, the
member stars lie in the region below the birthline
Steven W. Stahler and above the main sequence. In the very youn-
Department of Astronomy, University of gest clusters, most stars are crowded close to the
California, Berkeley, CA, USA birthline. In older groups, stars of lower mass lie
further below the envelope, while more massive
objects are already on the main sequence.
Keywords To see why there is a birthline, one needs to
consider stellar evolution prior to the pre-main-
Stellar evolution; Star formation sequence phase. The very youngest stars, known
326 Bitumen
as protostars, are optically invisible and still

acquire their masses from the surrounding Bitumen
clouds. As a protostar’s mass increases, so
does its radius. Indeed, the radius of the proto- Emmanuelle J. Javaux
star can be predicted with fair accuracy from Palaeobiogeology-Palaeobotany-
knowledge of its mass alone. Once cloud infall Palaeopalynology, Geology Department,
ceases, the star appears as a visible object on the Université de Liège, Liège, Belgium
appropriate pre-main-sequence track. The pro-
tostar mass-radius relationship dictates the ini-
tial positions of stars on their tracks. The locus Definition
of initial positions for various stellar masses
defines a curve in the HR diagram that is the Bitumen is the fraction of fossil organic matter in
birthline. sedimentary rocks that is soluble in organic sol-
An interesting feature of the birthline is that it vents (such as dichloromethane, hexanes, and
intersects the main sequence. Specifically, the methanol). It is a brown to black, highly viscous
two curves join at about ten solar masses. More material that includes diverse organic macromol-
massive objects have no pre-main-sequence ecules and chemical biomarkers. Bitumen is a
phase, but ignite hydrogen while they are still component in petroleum.
protostars. Thus, the early evolution of massive
stars is quite different from that of the more
common objects akin to the Sun. The intersection
See Also
of the birthline and main sequence can be seen
clearly in the HR diagram of the Orion Nebula
▶ Hydrocarbons
Cluster, the nearest region currently forming
▶ Kerogen
massive stars.
See Also
Black Hole
▶ Main Sequence, Star
▶ Pre-Main-Sequence Star Nikos Prantzos
▶ Protostars Institut d’Astrophysique de Paris, Paris, France
▶ Stellar Cluster
▶ T Association
Definition
A black hole is a region of space-time inside a

fictitious surface (the event horizon), from which
Hillenbrand L (1997) On the stellar population and star- nothing – not even light – may escape. Astro-
forming history of the Orion nebula cluster. Astron physical black holes are either stellar or
J 113:1733–1768 galactic. The former are the end products of the
Stahler SW (1983) The birthline for low-mass stars.
evolution of massive stars, and they are produced
Astrophys J 274:822–829
Stahler SW, Palla F (2004) The formation of stars. Wiley- through gravitational collapse (after the exhaus-
VCH, Weinheim, Chapter 16 tion of all nuclear fuels and the subsequent
Black Smoker, Organic Chemistry 327
▶ supernova explosion) if the mass of the residue the flanks of island arcs; their fossilized equiva-
is above the Oppenheimer-Volkoff limit lents are found in Phanerozoic and Precambrian
(2–3 MJ). Galactic (supermassive) black greenstone belts. The hydrothermal fluids form as
holes are found in the centers of most galaxies seawater seeps down into the oceanic crust where
(including the Milky Way), they have masses in it is heated and interacts with mainly basaltic B
the 106–109 MJ range, but their origin is poorly rocks. The heated fluid then ascends through
understood at present. fractures and exits at the sea floor.
See Also
See Also
▶ Supernova
▶ Hydrothermal Vent Origin of Life Models
▶ Mid-ocean Ridges
Black Smoker
Yves Fouquet Black Smoker, Organic Chemistry

Institut Français de Recherche pour l’Exploitation
de la mer (IFREMER), Issy-les-Moulineaux, Koichiro Matsuno
France Nagaoka University of Technology, Nagaoka,
Japan
Definition
Keywords
A black smoker is a sulfide-rich hydrothermal
vent or hot spring on the ocean floor. Hydrother- Chimney; Hydrothermal vents; Pyrite; Reducing
mal aqueous fluids with temperatures up to agents; Sulfide
400 C are issued from the vent. These fluids
contain high concentrations of dissolved metals,
sulfur species, and silica. Cooling and dilution as Synonyms
the fluid mixes with seawater causes the precipi-
tation of Fe-Cu-Zn-Pb sulfides, which give rise to Submarine hot spring
the dark color of the fluid (hence the name black
smoker). Accumulation of sulfides around the
vent builds up chimneys that may reach 60 m in Definition
height and support an exotic ecosystem of anaer-
obic fauna and ▶ chemolithoautotroph commu- A specific organic chemistry proceeds in the
nities. Long-lived black smoker complexes may vicinity of a black smoker, or a hydrothermal
accumulate large volumes of sulfide which, when vent, from which a cloud of black particles is
accreted to the continent, can be mined as ore emitted on the floor of the ocean. The main ingre-
deposits. Black smokers occur along modern dients of the black particles are metal sulfides.
▶ mid-ocean ridges, in back-arc basins and on Pyrite deposited around the smoker can serve as
328 Blackbody
an efficient reducing agent for synthesizing synthesize saturated fatty acids in the vicinity of
reduced carbon compounds from primary carbon hydrothermal vents, perhaps using cobalt-based
sources, such as carbon monoxide. catalysts where deleterious influences from sulfur
catalyst poisoning could be avoided. Organic
chemistry proceeding in and around black
Overview smokers could be quite rich, although some of
the synthetic reactions might mutually be inhibi-
A black smoker is a type of hydrothermal vent tive. To date, the only unambiguously abiogenic
found on the seafloor, appearing in the form of a organic compounds detected in these systems on
chimney-like structure. Hydrothermal vents are Earth are simple hydrocarbons such as methane,
located around fissures at mid-ocean ridges ethane, and propane.
where tectonic plates are moving apart, from
which heated water issues. Besides Earth, plane-
tary bodies that might possibly have or had such See Also
hydrothermal systems include Europa, a satellite
of Jupiter, and ancient Mars. ▶ Black Smoker
The cloud of black materials emitted out of the ▶ Fischer-Tropsch-Type Reaction
chimney includes a significant amount of black ▶ Hydrocarbons
particles made of metal sulfides. When the hot ▶ Hydrothermal Environments
water from the chimney comes in contact with the ▶ Hydrothermal Reaction
surrounding cold water, the black materials pre- ▶ Hydrothermal Vent Origin of Life Models
cipitate on top of the chimney, and it can gradu- ▶ Pyrite
ally grow in height until its inevitable collapse.
The piling up of the black chimney-like struc-
tures is considered to be a major source of sulfide
ore deposits on Earth. Huber C, Wächtershäuser G (1997) Activated acetic acid
▶ Pyrite is a typical metal sulfide deposited by by carbon fixation on (Fe, Ni)S under primordial con-
black smokers. It is a crystalline mineral com- ditions. Science 276:245–247
posed of iron and sulfur. Pyrite possesses a pos-
itive surface charge at ambient seawater pH and
accordingly can adsorb anions or organic com-
pounds having anionic carboxylate or phosphate
groups. In particular, when hydrogen sulfide Blackbody
reacts with iron in solution to form iron sulfide
under conditions expected in the vicinity of the Daniel Rouan
black smokers, the reaction produces electrons LESIA, Observatoire Paris-Site de Meudon,
that can reduce various chemicals if relevant Meudon, France
reactants are available. Iron sulfide formed in
the hydrothermal environment can serve as a
reducing agent that can drive a series of various Keywords
synthetic reactions. An experimental demonstra-
tion of the reducing capability associated with the Electromagnetic radiation; Temperature;
formation of pyrite from hydrogen sulfide and Emission; Spectrum
iron is the synthesis of acetate from carbon mon-
oxide (Huber and Wächtershäuser 1997).
It might therefore be expected that ▶ Fischer- Synonyms
Tropsch-type reactions using acetic acid or
formic acid as the primary carbon sources may Thermal emission
Blackbody 329
Definition most perfect astrophysical blackbody is without

any contest the Universe itself, which emits radi-
A blackbody is a hypothetical object that is a ation peaking in the millimetric range, the cosmic
perfect radiator, absorbing all radiation that microwave background (CMB); it fits Planck’s
impinges upon it and emitting over the whole law (at T = 2.73 K) to within 1 part in 1,000, as B
electromagnetic spectrum, however preferen- revealed by the satellite COBE. The blackbody
tially around a certain wavelength. The overall emission is indeed universal and is observed in a
power and the spectral energy distribution large variety of celestial objects at very different
depend only upon the object’s temperature. wavelengths. To give several examples, the emis-
sion of a star as the Sun, with T = 6,000 K, peaks
at l(mm) = 0.5 mm (visible); a planet as the
Overview Earth, with T = 300 K, emits mainly at
l(mm) = 10 mm (infrared); and the hot gas in a
At the laboratory, a good approximation of a cluster of galaxies, with T = 30 106 K, peaks
blackbody is a small-hole entrance to a cavity at l(mm) = 104 mm, i.e., in the X-ray domain.
with absorbing walls within a block maintained Obviously celestial objects are not strictly black
at a fixed temperature. The spectral distribution in general; this is why an approximation called
of the blackbody emission is described by the gray-body radiation is often used in models.
Planck’s law and depends only on the tempera-
ture of the object: it peaks at a wavelength
given by Wien’s law, approximately
lðmmÞ ¼ 3, 000=T ðKÞ. On the short-wavelength See Also
side, the intensity decreases very fast, while it
varies in a smoother way at wavelengths beyond ▶ Bremsstrahlung Radiation
the maximum (Fig. 1). The total power radiated ▶ Effective Temperature
by a blackbody is proportional to the fourth ▶ Electromagnetic Radiation
power of the temperature (Stephan’s law). As a ▶ Electromagnetic Spectrum
first approximation, stars’ and planets’ emission ▶ Grey Body
can be described by blackbody radiation. The ▶ Radiative Processes
Blackbody, Fig. 1 The COSMIC MICROWAVE BACKGROUND SPECTRUM FROM COBE

spectral distribution of the 1.2
Intensity, 10−4 ergs/cm2 sr sec cm−1
blackbody emission of the THEORY AND OBSERVATION AGREE

Universe as seen by the 1.0
satellite COBE. One notes
the strong asymmetry
between the short- and the 0.8
long-wavelength sides with
an abrupt exponential 0.6
decrease in the first case
and a smoother power-law
decrease in the second case 0.4
0.2
0.0
0 5 10 15 20
Waves/centimeter
330 Blue-Green Algae
Kroemer H, Kittel C (1980) Thermal physics, 2nd edn. ▶ UK Space Agency

W. H. Freeman Company, Los Angeles
Blue-Green Algae Bolometer
Daniel Rouan
▶ Cyanobacteria, Diversity and Evolution of
LESIA, Observatoire Paris-Site de Meudon,
Meudon, France
Blue-Green Bacteria Synonyms
▶ Cyanobacteria Pyrometer; Radiometer
Definition
BNSC
A bolometer is a detector that measures electro-
Michel Viso magnetic power incident on its absorbing surface
CNES/DSP/SME, Vétérinaire/DVM, Astro/ by sensing a temperature change through some
Exobiology, Paris Cedex 1, France temperature-dependent physical mechanism in
the detector, generally its electrical resistance.
In a typical bolometer, the radiative power is
Synonyms thermalized either by a crystal lattice or by the
electrons in the absorber. The small resulting
British National Space Council temperature rise is sensed by a thermometer – a
temperature-dependent semiconductor in the
case of a semiconducting bolometer or a
Definition superconducting metal film in the case of a
transition-edge superconducting detector. The
The British National Space Council (BNSC) was lower the temperature, the better the perfor-
established in 1985 replacing the Space Research mances of a bolometer: 100 mK is current in the
Management Unit that was running the UK space most sensitive instruments.
activities since 1963. The BNSC aimed to help Because they absorb any radiation, bolometers
Britain to get an optimal return of national and are able to measure the power received from
international activities in space, both economi- celestial objects over a broad range of wave-
cally and scientifically. BNSC was a voluntary length; they are however used mainly in the infra-
partnership between ten government departments red and millimeter bands.
and research councils. Their combined expendi-
ture on civil space amounts to around £180 mil-
lion per year.
The ▶ UK Space Agency took over in March See Also
2010 to become a full agency replacing the
BNSC by April 1, 2011. ▶ Bolometric Magnitude
Boron Isotopes 331
(Klochko et al. 2006). The two naturally occurring

Bolometric Magnitude and stable isotopes are 11B (80.1 %) and 10B
(19.9 %). The mass difference results in a wide
Daniel Rouan range of boron isotopic compositions in terrestrial
LESIA, Observatoire Paris-Site de Meudon, environments, and boron isotopic compositions in B
Meudon, France marine carbonates have been used as a paleo-pH
proxy of the oceans (Hemming and Hanson 1992).
Borates have also been found to stabilize ▶ ribose
Definition (Ricardo et al. 2004), which may have been impor-
tant for the origin of life on early Earth, although
The bolometric magnitude is a way to quantify the prebiotic availability of borate minerals has
the total power emitted by a celestial object in the been questioned (Hazen et al. 2008).
form of electromagnetic radiation. The bolomet-
ric magnitude is the ▶ magnitude of a celestial
object based on its flux integrated over the whole See Also
electromagnetic spectrum. By convention, the
bolometric magnitude of the star Vega is zero. ▶ Formose Reaction
The bolometric correction, BC, is the difference ▶ Ribose
between the visual and the bolometric
magnitudes.
See Also Dickson AG (1990) Thermodynamics of the dissociation

of boric acid in synthetic seawater from 273.15 to
318.15 K. Deep-Sea Res 37:755–766
▶ Magnitude Hazen RH, Papineau D, Bleeker W, Downs RT, Ferry JM,
▶ Magnitude, Absolute McCoy TJ, Sverjensky DA, Yang H (2008) Mineral
evolution. Am Mineral 93:1693–1720
Hemming NG, Hanson GN (1992) Boron isotopic compo-
sition and concentration in modern marine carbonates.
Geochim Cosmochim Ac 56:537–543
Klochko K, Kaufman AJ, Yao W, Byrne RH, Tossell JA
Borate (2006) Experimental measurement of boron isotope
fractionation in seawater. Earth Planet Sci Lett
Kateryna Klochko 248:261–270
Carnegie Institution of Washington, Washington, Ricardo A, Carrigan MA, Olcott AN, Benner SA
(2004) Borate minerals stabilize ribose. Science
DC, USA 303:196
Definition
Boron Isotopes
Borate is a common oxoanion (BO33) found in
boron-containing minerals (e.g., boracite, borax, Marc Chaussidon
kernite, colemanite, etc.), some borosilicates (e.g., Institut de Physique du Globe de Paris (IPGP),
tourmaline), and in association with natural car- Paris, France
bonates. In natural aqueous environments, borates
exist in equilibrium with boric acid B(OH)3 and
borate anion B(OH)4 . The pH-dependent equi- Keywords
librium between these species in seawater
(pKa = 8.597, (Dickson 1990)) is accompanied Boron; CAIs; Chondrites; Isotopes; Nucleosyn-
by boron isotope fractionation of 27.2‰ thesis; pH; Seawater
332 Boron Isotopes
Definition This nucleosynthetic origin makes the isotopic

composition of B in the Solar system quite
Boron has two stable isotopes (10B and 11B), 11B intriguing. In the Earth and meteorites, the
11 10
being the most abundant since the average B/ B ratio (4.04; Palmer and Swihart 1996)
terrestrial 11B/10B isotope ratio is of 4.04. is enriched in 11B relative to GCR. Boron in
Strong variations of this ratio were present in diffuse interstellar clouds is also 11B-rich
the presolar molecular cloud and in the accretion (11B/10B = 3.4 0.7, Lambert et al. 1998) rela-
disk, part of them due to the radioactive decay of tive to GCR. This 11B enrichment has been pro-
10
Be to 10B. Low-temperature isotopic fraction- posed to result from the addition of B made by
ations are responsible of large (up to 100 ‰, low-energy spallation which would be enriched
where ‰ is parts per mil) variations of the in 11B according to predictions made from avail-
11 10
B/ B ratio in seawater and terrestrial rocks. able cross sections (Reeves 1994) and/or to the
Some of these variations may be used to recon- addition of B made by neutrino-induced spall-
struct paleo-seawater pH and paleo-atmospheric ation of C in type II ▶ supernovae (Lambert
PCO2. et al. 1998). These processes are likely to operate
at galactic time scales but irradiation processes
around the young forming Sun also produced a
Overview fraction of the Solar System boron. This is dem-
onstrated (McKeegan et al. 2000) by the presence
Boron is one member of the lithium-beryllium- in Ca-Al-rich refractory inclusions (▶ CAIs)
boron (Li-Be-B elements) clan, three elements from chondritic meteorites of 10B excesses due
which share some unique properties due to a com- to the radioactive decay of short-lived 10Be (10Be
mon nucleosynthetic history. The cosmic abun- b decays to 10B with a half-life of 1.39 Ma). The
dance (or solar abundance) of Li-Be-B elements magnitude of the10B excesses implies that the
(Lodders 2003) is a factor of 107 – 1010 lower 10
Be/9Be ratio at the time of formation of CAIs
than elements of neighboring atomic mass such as was up to 8.8 104 (Chaussidon et al. 2006)
He, C, and O. This is due to the fact that light but large variations are expected in this ratio
elements (D and Li-Be-B clan) cannot be synthe- depending on the conditions of the irradiation
sized by nuclear reactions in stellar interiors at processes in the early Solar System (Chaussidon
temperatures higher than 106 K, being rather and Gounelle 2006). One attractive model is the
destroyed at these temperatures (Burbidge formation of 10Be and of B isotopes (10B and 11B)
et al. 1957). Contrary to 7Li, for which a fraction by reactions between flares emitted by the young
is produced during the Big-Bang nucleosynthesis, active Sun when a classical T-Tauri star and
the two stable isotopes of B (10B and 11B) are grains and/or gas at the inner edge of the
produced in the interstellar medium by endothermic accretion disk.
spallation reactions. These reactions occur at high While B isotopic variations inherited from
energies, typically higher than a few MeV (Reeves nucleosynthesis might be present in the accretion
et al. 1970). High-energy collisions between accel- disk (Chaussidon and Robert 1995, 1998),
erated protons (and alpha particles) and O and they are most probably averaged at planetary
C nuclei at rest (mostly 16O and 12C) are responsible scale. However, B isotopic variations around
of the production of 10B and 11B with an isotope this ratio of 4.04 are widespread in terrestrial
ratio 11B/10B 2.5, as modeled from available rocks. They are due to low-temperature stable
cross sections (Meneguzzi et al. 1971) and as mea- isotopic fractionations and are generally reported
sured in present-day galactic cosmic rays (GCR) as deviations in permil from the 11B/10B ratio
(11B/10B from 1.5 to 3.2 depending on the of the international standard (NBS boric acid
energy, Krombel and Wiedenbeck 1988). The SRM 951 with 11B/10B = 4.04367, Catanzaro et al.
poor efficiency of these reactions is responsible of 1970) using the classical delta notation (d11Bsample =
the low cosmic abundance of B. ((11B/10B)sample/(11B/10B)SRM951 1) 1,000).
Boron Isotopes 333
Boron in the silicate Earth is grossly distrib- allows the boron isotopic fractionation to be cal-
uted as follows: 10 % in seawater, 35 % in culated between a mineral and the pH of the solu-
the crust (continental and oceanic), and 55 % tion and has been widely used and investigated to
in the mantle (upper plus lower mantle). Very try to reconstruct paleovariations of seawater pH
little is known for boron isotope variations in and atmospheric PCO2 (e.g., Pearson and Palmer B
organic matter (Williams et al. 2001) because 2000, refs. therein). These reconstructions are
of analytical difficulties, even though boron is complicated by the fact that the d11B of seawater
a key element for life and has a complex might have changed over geological times
biogeochemical cycle (Park and Schlesinger (Lemarchand et al. 2000) and that a significant
2002). Boron in the primitive mantle has a fraction of B3 can also be adsorbed. However,
d11B 10 ‰ as indicated by oceanic basalt boron isotopes are one of the best tools used to
glasses (Chaussidon and Marty 1995) and this infer the pH of a solution from which a mineral
is most probably the bulk d11B of the Earth. grew. This has been useful to unravel the origin of
There is no significant isotopic fractionation the so-called vital effect on the oxygen isotopic
associated with the extraction of boron from fractionation between biogenic carbonates and
the mantle to the crust, because boron isotopic seawater. Micrometer scale variations of d11B
fractionations at mantle temperatures are mini- values in coral skeletons have shown that strong
mum. This is confirmed by analyses of tourma- pH variations occurred during the formation of
lines from granitic rocks of various ages which calcite and that these changes of pH modify the
allow to define an average d11B for the continental kinetic of oxygen isotopic exchanges between
crust (at a mean age of 2 Ga) between 8 ‰ dissolved carbonate species (H2CO3, HCO3 and
and 13 ‰ (Chaussidon and Albarède 1992). CO32) and water via the two reactions of hydra-
However, in more details, mass balance implies tion and hydroxylation (Rollion-Bard et al. 2003).
that the d11B of the crust has evolved over geologic Boron isotopes in sedimentary rocks are thus key
times in response to crustal growth and to the for the reconstruction of past paleo-environmental
change of the relative fractions of B hosted in the changes on Earth.
main surface reservoirs. In fact, a major isotopic
fractionation occurs for boron during adsorption
See Also
processes on clay minerals (Schwartz et al. 1969;
Palmer et al. 1987) and various other phases such
▶ CAIs
as, for instance, humic acids (Lemarchand
▶ Chondrite
et al. 2005). This isotopic fractionation is due to
▶ Nucleosynthesis, Stellar
the facts that (1) boron in solution occurs both as
boric acid (B(OH)3, named B3 in the following)
and borate anion (B(OH)4, named B4 in the fol- References and Further Reading
lowing), the relative proportions of the two species
depending on the acidity constant (pK) of boric Burbidge EM, Burbidge GR, Fowler WA, Hoyle F (1957)
Synthesis of the elements in stars. Rev Mod Phys
acid, which itself depends on temperature 29:547–650
and salinity, and (2) there is a strong equilibrium Catanzaro EJ et al (1970) Standard reference materials:
isotopic fractionation between boric acid and boric acid; isotopic, and assay standard reference
borate anion (aB3-B4 = 0.97352 at 25 C, Pagani materials. National Bureau of Standards Special Pub-
lication 260-17. US Government Printing Office,
et al. 2005; Klochko et al. 2006; Rollion-Bard and Washington, DC, 70 pp
Erez 2010). Boron adsorbed at the surface of grow- Chaussidon M, Albarède F (1992) Secular boron isotope
ing minerals (clays, carbonates) is dominantly B4, variations in the continental crust: an ion microprobe
which is thus strongly isotopically fractionated study. Earth Planet Sci Lett 108:229–241
Chaussidon M, Gounelle M (2006) Irradiation processes
from the solution. During crystal growth, this in the early solar system. In: Lauretta DS, McSween
adsorbed boron is incorporated into the structure HY Jr (eds) Meteorites and the early solar system
of the minerals (Spivack et al. 1987). This property II. University of Arizona Press, Tucson, pp 323–339
334 Brazilian Space Agency
Chaussidon M, Marty B (1995) Primitive boron isotope Reeves H, Fowler W, Hoyle F (1970) Galactic cosmic rays
composition of the mantle. Science 269:383–386 origin of Li, Be and B in stars. Nature 226:727–729
Chaussidon M, Robert F (1995) Nucleosynthesis of 11B- Rollion-Bard C, Chaussidon M, France-Lanord C (2003)
rich boron in the pre-solar cloud recorded in meteoritic pH control on oxygen isotopic composition of symbi-
chondrules. Nature 374:337–339 otic corals. Earth Planet Sci Lett 215:275–288
Chaussidon M, Robert F (1998) 7Li/6Li and 11B/10B var- Rollion-Bard C, Erez J (2010) Intra-shell boron isotope
iations in chondrules from the Semarkona ratios in the symbiont-bearing benthic foraminiferan
unequilibrated chondrite. Earth Planet Sci Lett Amphistegina lobifera: Implications for d11B vital
164:577–589 effects and paleo-pH reconstructions. Geochim
Chaussidon M, Robert F, McKeegan KD (2006) Li and Cosmochim Acta 74:1530–1536
B isotopic variations in an Allende CAI: evidence for Schwartz HP, Agyei EK, McMullen CC (1969) Boron
the in situ decay of short-lived 10Be and for the possi- isotopic fractionation during clay adsorption from sea-
ble presence of the short-lived nuclide 7Be in the early water. Earth Planet Sci Lett 6:1–5
solar system. Geochim Cosmochim Acta 70:224–245 Spivack AJ, Palmer MR, Edmond JM (1987) The sedi-
Klochko K, Kaufman AJ, Yao W, Byrne RH, Tossell mentary cycle of boron isotopes. Geochim
J (2006) Experimental measurements of boron isotopic Cosmochim Acta 51:1939–1949
fractionation in seawater. Earth Planet Sci Lett Williams LB, Hervig RL, Wieser ME, Hutcheon I (2001)
248:276–285 The influence of organic matter on the boron isotope
Krombel KE, Wiedenbeck ME (1988) Isotopic composi- geochemistry of the gulf coast sedimentary basin. USA
tion of cosmic-ray boron and nitrogen. Astrophys Chem Geol 174:445–461
J 328:940–953
Lambert DL, Sheffer Y, Federman SR, Cardelli JA, Sofia
UJ, Knauth DC (1998) The 11B/10B ratio of local
interstellar diffuse clouds. Astrophys J 494:614–622
Lemarchand D, Gaillardet J, Lewin E, Allègre CJ Brazilian Space Agency
(2000) The influence of rivers on marine boron iso-
topes and implications for reconstructing past ocean
pH. Nature 408:951–954 ▶ AEB
Lemarchand E, Schott J, Gaillardet J (2005) Boron isoto-
pic fractionation related to boron sorption on humic
acid and the structure of surface complexes formed.
Lodders K (2003) Solar system abundances and conden- Breccia
sation temperatures of the elements. Astrophys
J 591:1220–1247 Daniele L. Pinti
McKeegan KD, Chaussidon M, Robert F (2000) Incorpo-
ration of short-lived 10Be in a calcium-aluminium-rich
inclusion from the Allende meteorite. Science Geodynamics, Université du Québec à Montréal,
289:1334–1337 Montréal, QC, Canada
Pagani M, Lemarchand D, Spivack A, Gaillardet J (2005)
A critical evaluation of the boron isotope-pH proxy:
the accuracy of ancient pH estimates. Geochim
Cosmochim Acta 69:953–961 Synonyms
Palmer MR, Spivack AJ, Edmond JM (1987) Temperature
and pH controls over isotopic fractionation during Rubblerock
adsorption of boron on marine clay. Geochim
Cosmochim Acta 51:2319–2323
Palmer MR, Swihart GH (1996) Boron isotope geochem-
istry: an overview. In: Grew ES, Anovitz LM (eds) Definition
Boron, mineralogy, petrology and geochemistry. Rev
mineral, vol 33. Mineralogical Society of America,
Breccia is a rock composed of angular fragments.
pp 709–744
Park H, Schlesinger WH (2002) Global biogeochemical It is produced by sedimentary, magmatic, or tec-
cycle of boron. Glob Biogeochem Cycles 16:1072 tonic processes. A sedimentary breccia is a
Pearson PN, Palmer MR (2000) Atmospheric carbon diox- coarse-grained (particle size >2 mm) rock com-
ide concentrations over the past 60 million years.
posed of angular rock fragments held together by
Nature 406:695–699
Reeves H (1994) On the origin of the light elements cement or dispersed in a fine-grained solidified
(Z < 6). Rev Mod Phys 66:193–216 matrix. It originates as a result of sedimentary
Brown Dwarf 335
processes such as talus accumulation analogy with ▶ blackbody radiation and is, as
(sedimentary breccia), disturbance during sedi- well, considered as thermal emission, as opposed
mentation (intraclastic breccia), or collapse of to nonthermal processes, like synchrotron emis-
rocky surfaces such as the roof of a cave sion. The notable difference with blackbody
(collapse breccia). Magmatic breccia results emission is that bremsstrahlung features a plateau B
from explosive eruptions (pyroclastic breccia), of quasi-constant intensity in a broad region of
hydrovolcanic fragmentation in diatremes wavelengths, while the blackbody emission
(funnel-shaped pipes), or mechanical fragmenta- exhibits a pronounced peak.
tion in intrusive settings. Tectonic processes such
as frictional slip along a fault produce fault brec-
cia. Breccia containing angular clasts and fine- See Also
grained matrix produced by meteoritic impact is a
common rock on planetary surfaces. The ▶ anor- ▶ Blackbody
thosite breccia of the lunar highlands is a well- ▶ Radiative Processes
known example. ▶ Radio Astronomy
See Also
Brethren of Purity
▶ Crater, Impact
▶ Impactite ▶ Ikhwan al-Safa
▶ Moon, The
▶ Suevite
British National Space Council
▶ BNSC
Bremsstrahlung Radiation
Daniel Rouan
Meudon, France
Broadband Line Survey
▶ Molecular Line Survey

Synonym
Free-free emission
Brown Dwarf
Definition Gibor Basri

Astronomy Department, MC 3411, University of
Bremsstrahlung is a German word used in phys- California, Berkeley, CA, USA
ics to describe the electromagnetic radiation
emitted by a plasma in which high-speed charged
particles (electrons, protons) are accelerated by Keywords
other charged particles (ions). The bremsstrah-
lung radiation emitted by a plasma shares some Very cool stars; Very low-mass stars
336 Brown Dwarf
Synonyms pressure. This form of pressure does not depend

on temperature, but is produced by the fact that
Substellar objects electrons cannot occupy the same low-energy
quantum states when forced too close to each
other. This produces the counterintuitive prop-
Definition erty that adding more mass to the object means
a higher density core is needed to support it, so
A brown dwarf is an astronomical object inter- the object actually becomes smaller. Thus, brown
mediate in mass between stars and planets, some- dwarfs are generally a little smaller in size than
times called a substellar object. Brown dwarfs are planets like Jupiter, and the most massive ones
different from stars in that they never achieve are the smallest (and thereby most dense). Since
stable luminosities by nuclear fusion and differ- objects that degenerate are no longer generating
ent from planets in that they are capable of energy, they are on an endless slide to cooler,
nuclear fusion early in their evolution. Their pri- fainter states (Chabrier and Baraffe 2000;
mary means of pressure support is electron Burrows et al. 2001). It is worth noting that stars
degeneracy. When very young, they closely like the Sun also end in a similar phase after
resemble very low-mass stars, but later they fusion ceases; they are then known as “white
cool below the minimum stellar temperature and dwarfs” (which are much more massive and so
develop mineral grains in their atmospheres. can be as small as the Earth and initially much
hotter – hence “white”).
Overview
Basic Methodology
Brown dwarfs are considered by astronomers to
be substellar objects, meaning they are interme- Brown dwarfs were first hypothesized in 1963 by
diate in mass between a planet and a star. That is, Shiv Kumar, who called them “black” dwarfs. Jill
their masses are above that of the most massive Tarter proposed the name “brown dwarf” in 1975.
planet (13 Jupiter masses) but below the least They are not actually brown, but would appear to
massive star (75 Jupiter masses or 0.075 solar the eye from deep red to magenta depending on
masses). These objects contract under the influ- their temperature. No brown dwarfs were defini-
ence of self-gravity. While doing so, they fuse tively known before 1994. In order to distinguish
deuterium (“heavy hydrogen,” a rare isotope pro- (brighter, warmer) brown dwarfs from stars of the
duced in the Big Bang) during their first few same temperature, one can search for lithium in
million years of life. Deuterium fusion almost their spectrum (the primoridal complement of lith-
entirely halts the contraction while it is going ium created by the Big Bang is destroyed when
on. At the upper planetary mass boundary, deu- hydrogen fusion begins). Objects as cool as red or
terium fusion cannot occur because the central brown dwarfs are fully convective, so all parts of
core never becomes hot and dense enough. True the object are mixed to the core on fairly short
stars also contract past the point of deuterium timescales, and the core properties are simply
fusion, but their central temperature then climbs related to mass and age. Brown dwarfs below
until they fuse ordinary hydrogen, which stabi- 60 Jupiter masses will never destroy lithium,
lizes the star and allows it to join the “main while all low-mass stars will, within a few tens
sequence.” Technically, brown dwarfs above of millions of years. Alternatively, one can look
60 Jupiter masses also begin to fuse ordinary for (much fainter) objects below the minimum
hydrogen but then stabilize and the fusion stops. stellar temperature (which was tried first with stel-
Brown dwarfs stabilize against contraction lar companions). In 1995, both methods produced
because their cores are dense enough to hold verified discoveries (Basri 2000). The lithium test
themselves up with “free electron degeneracy” was successfully employed at UC Berkeley on an
Brown Dwarf 337
object in the Pleiades (PPl 15, which later also VO at visible wavelengths, whereas brown
proved to be the first brown dwarf binary). In the dwarfs have fairly smooth spectra (dust has
same time frame, astronomers at Palomar Obser- replaced the molecules) and enormous features
vatory and Johns Hopkins University found a com- due to atomic transitions of alkali elements, espe-
panion to a red dwarf star (Gliese 229; “red cially sodium and potassium (and lithium), which B
dwarfs” are very low-mass stars). The detection form above the dust. In the infrared, brown
of methane in the companion’s spectrum showed dwarfs show molecular features due to water,
that it has a surface temperature less than carbon monoxide, and in the cooler ones, meth-
1,300 K. Its extremely low luminosity, coupled ane. These spectral differences led to the delin-
with the age of its stellar companion, implies that eation of the first new astronomical spectral class
it has about 50 Jupiter masses. Christened Gliese (“L”) to be defined since the early twentieth cen-
229 B, this was the first object widely accepted as a tury. The coolest actual stars also extend into this
brown dwarf. L class. The next new spectral class, “T,” is
defined by the appearance of methane in the
infrared spectra of brown dwarfs (Kirkpatrick
Key Research Findings 2005) with surface temperatures below about
1,300 K.
The surface temperatures of brown dwarfs The dust can form clouds, which gives rise to a
depend on both their mass and age; they start sort of “weather” that can sometimes be detected
off as large and hot as they will ever be. A youn- as rotational modulation of brightness variations
ger, less massive object can have the same tem- (clouds will be darker and cooler). For objects at
perature as an older, more massive object. This the boundary between spectral classes L and T,
makes them fundamentally different from main the refractory clouds sink down below the atmo-
sequence stars, which are hotter when more mas- sphere. The colors and spectra of very cool
sive. The most massive and youngest brown objects are complicated by the formation of
dwarfs have temperatures as high as 2,800 K clouds, and the most successful models have to
(overlapping with red dwarfs). All brown dwarfs incorporate various amounts of mixing of cloudy
eventually cool below the minimum main and clear atmospheres (Saumon and Marley
sequence stellar temperature of about 2008). The spectral sequence has a looser relation
1,800 K. The oldest and least massive can be as to the temperature sequence in this regime. The
cool as about 500 K. Since age and size are often actual formation of condensates, their sizes and
hard to determine, the mass of a brown dwarf will settling properties, are not well understood (even
be uncertain unless it is located within a cluster the formation of clouds on the Earth is tricky).
(or binary) of known age or is in a known orbit. It appears that brown dwarfs form in much the
Eclipsing binary brown dwarf systems (where same way as stars, constituting the very low-mass
both size and mass can be independently found) end of the star formation process (Luhman
are extremely valuable in testing models for et al. 2007). They possess circumstellar disks
brown dwarfs (the first systems have recently when forming and thus might also sometimes
been discovered). have planets (a couple of giant planet compan-
Objects below about 2,200 K have mineral ions have been found already). They exhibit all
grains in their atmospheres and hence are the phenomenology of newly forming stars, but
“dusty” (Allard et al. 2001; Lodders and Fegley with lower accretion rates. Brown dwarfs, like Gl
2006). The dust is made of the most refractory 229B, the first one discovered, can be found as
compounds: Ca, Al, and Ti oxides like corundum companions to stars or even to each other. Their
(Al2O3) and perovskite (CaTiO3). The presence binary fraction is comparable to that for red
of dust makes the spectra of most brown dwarfs dwarfs (about one quarter) and in line with the
look quite different than red dwarfs; the latter decrease observed with stellar mass along the
show many bands from molecules like TiO and main sequence. An alternative formation
338 Brownlee Particle
scenario, that brown dwarfs are stars whose birth ▶ Planet

is interrupted by being ejected from a multiple ▶ Protostars
system before they can gather enough mass, is not ▶ Spectral Type
supported by most observations. The disk life- ▶ Star Formation, Theory
times are not very different, and the wide binary
frequency is not easily compatible with that
hypothesis. There also seems to be a population
of isolated objects forming like stars all the way Allard F, Hauschildt HP, Alexander RD, Tamanai A,
down into the high planetary mass domain. Schweitzer A (2001) The limiting effects of dust in
One respect in which brown dwarfs are more brown dwarf model atmospheres. Astrophys
like giant planets than stars is that they tend to J 556:357–372
Basri G (2000) Annu Rev Astron Astrophys 38:485–519
rotate in a few hours rather than days or weeks Burrows A, Hubbard WB, Lunine JI, Liebert J (2001) Rev
like most stars. This has to do with the fact that Mod Phys 73:719–765
although young brown dwarfs are magnetically Chabrier G, Baraffe I (2000) Annu Rev Astron Astrophys
active (as are young stars), they are not able to 38:337–377
Kirkpatrick J (2005) Davy. Annu Rev Astron Astrophys
sustain magnetically driven mass loss (which also 43:195–245
causes angular momentum losses that slow the Lodders K, Fegley B Jr (2006) Chemistry of low mass
rotation down in stars). The reason behind this is substellar objects. In: Mason JW (ed) Astrophysics
that as their surfaces cool, the magnetic field gets update, vol 2. Springer, Heidelberg, p 1
Luhman KL, Joergens V, Lada C, Muzerolle J, Pascucci I,
decoupled from the increasingly neutral plasma, White R (2007) The formation of brown dwarfs: obser-
making it difficult to sustain the coronal heating vations. In: Reipurth B, Jewitt D, Keil K (eds) Pro-
that drives winds (Reiners and Basri 2008). tostars and planets V. University of Arizona Press,
Infrared sky surveys and other techniques Tucson, pp 443–457
Reiners A, Basri G (2008) Astrophys J 684:1390–1403
have now uncovered hundreds of brown dwarfs. Saumon D, Marley MS (2008) Astrophys J 689:
By studying very young clusters, we are able to 1327–1344
assess how many brown dwarfs form relative to
stars. The result is that there seem to be as many
as a few percent of brown dwarfs compared to
stars, which implies there are billions of them in
our Galaxy. Before their discovery, it was Brownlee Particle
thought that brown dwarfs might be a significant
component of “dark matter” (they are too faint to ▶ Interplanetary Dust Particle
be seen except when very close by), but they do
not add up to nearly enough mass to be interesting
in that respect. From a physical point of view,
each brown dwarf begins with stellar surface Bruno, Giordano
temperatures and ends with surface temperatures
like those of young giant planets by the time the Fernando B. Figueiredo
current age of the universe has passed. We have CITEUC, University of Coimbra, Coimbra,
much to learn from studying them. Portugal
See Also History
▶ Binary Stars, Young Giordano Bruno (1548–1600) was Italian hetero-

▶ Dwarf Star dox philosopher and a theologian. Because of his
▶ Low Mass Star ideas and writings, Giordano Bruno left Naples,
▶ Main Sequence, Star his home town, in 1576. In 1583, he abandoned
B€
ucherer-Bergs Synthesis 339
his Dominican habit and left Italy, living thereaf-

ter in France, England, and Germany. € cherer-Bergs Synthesis
Bu
A supporter of Copernicus’s heliocentric the-
ory, of which he became a staunch supporter, Robert Pascal
Giordano Bruno collected countless enemies for Institut des Biomolécules Max Mousseron B
their unorthodox ideas. In his book De l’Infinito, CC1706, Université de Montpellier II,
Universo e Mondi (1584), he argued that the Montpellier, France
universe was infinite, that it contained an infinite
number of worlds, and that these are all inhabited
by intelligent beings.
In 1591, he left to Venice, where in 1592 he Synonyms
got arrested and tried by the Inquisition. After he
recanted, Bruno was sent to Rome, in 1592, for Hydantoin formation
another trial. After 8 years of successive interro-
gations, he was declared a heretic. In 1660, Bruno
was burned at the Inquisitorial stake because of
his Copernicanism and his belief in the infinity of Definition
inhabited worlds.
The B€ucherer-Bergs synthesis (Ware 1950) of
▶ hydantoins is carried out by reacting cyano-
hydrins (the addition products of cyanide anion
BSL to aldehydes or ketones) with ammonium car-
bonate (or CO2 and NH3). Its mechanism,
▶ Biological Safety Level closely related to the ▶ Strecker synthesis,
€ cherer-Bergs
Bu
Synthesis, Fig. 1
340 Buckminsterfullerene
involves a-aminonitriles as intermediates that

are processed via a carbon dioxide-promoted Buffer
reaction (Fig. 1) (Pascal et al. 2005).
The Hydantoins can be ring-opened Juli Peretó
hydrolytically to form N-carbamoylamino acids. Institut Cavanilles de Biodiversitat i Biologia
It is considered relevant to amino acid formation Evolutiva, Universitat de València, València,
on the early Earth because it is likely to have Spain
prevailed over Strecker synthesis in a CO2-rich
atmosphere. Peptides are potentially produced
through conversion of N-carbamoylamino acids Definition
to amino acid N-carboxyanhydrides (Pascal
et al. 2005) (Fig. 1). Buffer is a solution of a conjugated pair of acid
and base that at a pH around its pKa can tolerate
moderate additions of acid or base without a
remarkable pH change.
See Also
▶ Amino Acid N-Carboxy Anhydride

▶ N-Carbamoyl Amino Acid Buffon’s Conception of Origins
▶ Strecker Synthesis of Life
Stéphane Tirard
Centre François Viète d’Histoire des Sciences
References and Further Reading et des Techniques EA 1161, Faculté des
Sciences et des Techniques de Nantes, Nantes,
Pascal R, Boiteau L, Commeyras A (2005) From the France
prebiotic synthesis of a-amino acids towards a primi-
tive translation apparatus for the synthesis of peptides.
Top Curr Chem 259:69–122
Ware E (1950) The chemistry of hydantoin. Chem Rev Keywords
46:403–470
History of earth; Spontaneous generation
History
Buckminsterfullerene
Georges-Louis Leclerc, Comte de Buffon
(1707–1788) was one of the most important
▶ Fullerene
French naturalists of the eighteenth century. In
1739, he became Intendant du Jardin et du Cab-
inet d’ Histoire Naturelle du Roi. For four
decades (1749–1789), he published his Histoire
Naturelle (22 volumes, with the supplements).
Buckyball From the beginning of this voluminous work,
he presented several important concepts and used
▶ Fullerene them in his description of living beings.
Bulk Silicate Earth 341
According to him, every animal and vegetable

organism was made of “organic molecules,” Building Blocks of Primitive Life
which were the smallest living entities. In each
body, the organic molecules took the mark of the ▶ Endogenous Synthesis
“internal mold” specific for each species. During B
reproduction, the male and the female would give
some organic molecules with the mark of the
parental internal molds, and the young organism
would then depend on it. Bulk Silicate Earth
It is very important to note that Buffon was not
a transformist. Indeed, he conceived some mod- Daniele L. Pinti
ifications in the limit of each species, which could GEOTOP Research Center for Geochemistry and
be reversible; however, he never accepted any Geodynamics, Université du Québec à Montréal,
link between species by means of modifications. Montréal, QC, Canada
As far as the history of Earth is concerned,
Buffon’s conception evolved during his career.
In the first volume of his Histoire Naturelle, he
developed a cyclic conception of the history of Synonyms
Earth. Thirty years later, he claimed a sagittal
theory and gave a description of “the Epochs of Primitive mantle composition (Acronyms: BSE)
the Nature” in a supplementary volume (V) of his
Histoire Naturelle. He described seven epochs.
At the beginning, the Earth was an incandescent
sphere of matter. The temperature progressively Definition
decreased and the Earth became solid. Therefore,
his history of Earth was irreversible and Bulk Silicate Earth (BSE) refers to the original
depended on the decrease in temperature. chemical composition of the silicate part of the
Buffon asserted that species were formed by Earth after the accretion and separation of a core
spontaneous generation and had been constituted but prior to differentiation of the first crust. For
by the organic molecules that were very abundant this reason, BSE is comparable to the composi-
in the soil during the first epochs of nature. For tion of the Earth’s “primitive mantle.” There are
him, spontaneous generations were still active in several models to evaluate the BSE initial com-
nature, but only for the little species. position. The classic model is to infer BSE from
the chemical composition of the most primitive,
undifferentiated meteorites, i.e., the CI-class
carbonaceous chondrites. Using this model,
See Also refractory lithophile elements (i.e., those which
did not enter the core but remained in the silicate
▶ Spontaneous Generation, History of mantle) are given relative abundances as they
occur in chondrites. Other models point to dif-
ferences in ratios of 142Nd/144Nd and 3He/4He
References and Further Reading between primitive, mantle-derived lavas and
chondrites to derive a somewhat different,
Roger J (1993) Les sciences de la vie dans la pensée
française au XVIIIe siècle (nouvelle édition). Albin nonchondritic composition of the primitive
Michel, Paris mantle.
342 Burgess Shale Biota
See Also Overview
▶ Archean Mantle The Burgess Shale biota figures strongly in dis-

▶ Carbonaceous Chondrite cussions of early animal evolution and the Cam-
▶ Hadean Mantle brian explosion of life. Unlike most fossil
▶ Radiogenic Isotopes localities, the Burgess Shale yields exquisitely
▶ Silicon Isotopes preserved remains of comparatively soft-bodied
animals, alongside more familiar and routinely
preserved biomineralizing (“shelly”) groups,
thus revealing a far more complete picture of
contemporaneous diversity, disparity, and eco-
Burgess Shale Biota logic range.
Interpreting the Burgess Shale animals has not
Thomas H. P. Harvey been straightforward, not least because they are
Department of Earth Sciences, University of separated from living relatives by approximately
Cambridge, Cambridge, UK half a billion years of evolution. Following the
early work of Walcott, the fossils were generally
assigned to members of familiar, living animal
Keywords groups. Later research questioned this approach,
which has been described by Gould as
Cambrian; Evolution; Metazoans; Paleobiology “shoehorning,” because the unusual morphologies
of many of the fossils apparently excluded them
from known groups. This led to the view that the
Definition Cambrian Period was a time of unprecedented
evolutionary experimentation, with the invention
The Burgess Shale biota is an assemblage of of many phylum-level body plans, only some of
exceptionally preserved fossil organisms from which survived to the present day. Increasingly,
rock units within the Burgess Shale Formation however, this conclusion is viewed as an artifact of
of the Rocky Mountains of British Columbia, classification. The application of cladistic methods
Canada. It is dated at ca. 505–510 Ma, within of phylogenetic analysis suggests that the Burgess
the middle part of the Cambrian Period of geo- Shale taxa, like all other fossils, can be accommo-
logic time, and is renowned for providing a dated in “stem groups,” which possess some but
detailed snapshot of early animal evolution. not all of the characters shared by extant members
of a lineage (the “crown”). In this light, the Bur-
gess Shale biota helps to resolve the evolutionary
History steps involved in the emergence of the various
animal body plans.
The principal fossil locality near Mount Bur- The exceptional fossils of the Burgess Shale are
gess was discovered in 1909 by Charles preserved as macroscopic carbonaceous compres-
D. Walcott, Secretary of the Smithsonian Insti- sions in fine-grained rocks. Cambrian fossil
tution. He quickly recognized the importance of assemblages of comparable “Burgess-Shale-
the fossils and coined the term “Burgess Shale.” type” preservation have subsequently been identi-
Later, expeditions from Harvard University, fied from some tens of sites around the world,
the Geological Survey of Canada, and the among the most spectacular being the Sirius Passet
Royal Ontario Museum discovered new locali- biota of northern Greenland and the Chengjiang
ties and amassed substantial new collections of biota of Yunnan Province, China, both several
fossils. million years older than the Burgess Shale itself.
Butadiynyl Radical 343
The preservational window that provides this par- envelopes of evolved carbon stars (evolved stars
ticular view onto early and middle Cambrian life whose atmospheres contain more carbon than
appears to be restricted in time and to particular oxygen, the excess carbon presumably produced
marine settings. The origins and evolutionary fate by helium fusion in the latter stages of the star’s
of Burgess-Shale-type biota are thus unclear and life). It is an intermediary in the rich carbon B
form a focus of current research. chemistry of these stars and of cold, dark inter-
stellar clouds; both contain polyacetylenes,
▶ cyanopolyynes, and related carbon-chain spe-
See Also cies. Both deuterated and 13C variants of C4H
have been detected in astronomical sources
▶ Cambrian Explosion (Cernicharo et al. 2000).
▶ Chengjiang Biota, China
History
Briggs DEG, Erwin DH, Collier FJ (1994) The fossils of The astronomical identification of C4H in the
the Burgess Shale. Smithsonian Books, Washington, envelope of the carbon star IRC + 10216
DC (Guélin et al. 1978) prior to laboratory measure-
Conway Morris S (1998) The crucible of creation: the ments of the rotational spectrum provides another
Burgess Shale and the rise of animals. Oxford Univer-
sity Press, Oxford example of the ability of high-frequency radio
Gould SJ (1989) Wonderful life: the Burgess Shale and the astronomy to contribute to fundamental molecu-
nature of history. Norton, New York lar physics (cf. the discussion for ▶ C3N). Subse-
quently, this radical was also detected in
interstellar molecular clouds (Irvine et al. 1981).
Butadiyne
See Also
▶ Diacetylene
▶ Cyanoethynyl Radical
▶ Cyanopolyyne
▶ Deuterium
▶ Molecular Cloud
Butadiynyl Radical ▶ Molecules in Space
▶ Radical
William M. Irvine

Synonyms
Cernicharo J, Guélin M, Kahane C (2000) A l 2 mm
molecular line survey of the C-star envelope IRC +
C4H 10216. Astrophys J Suppl 142:181–215
Guélin M, Green S, Thaddeus P (1978) Detection of the
C4H radical toward IRC +10216. Astrophys J 224:
Definition L27–L30
Irvine WM, Höglund B, Friberg P, Askne J, Elldér
J (1981) The increasing chemical complexity of the
The univalent five-atom ▶ radical C4H is found taurus dark clouds – detection of CH3CCH and
both in interstellar ▶ molecular clouds and in the C4H. Astrophys J 248:L113–L117
344 Butanedioic Acid
Butanedioic Acid Butyrine
▶ Succinic Acid ▶ Aminobutyric Acid
Butanenitrile Butyronitrile
▶ Propyl Cyanide ▶ Propyl Cyanide
Butlerow Reaction
C
C2H Definition
▶ Ethynyl Radical The cation C3H+ is thought to play an important

role in gas phase reactions in the ▶ interstellar
medium that lead to the formation of small hydro-
C2H4O carbons. The neutral radical, C3 H
(▶ propynylidyne) can exist in both a linear
▶ Ethylene Oxide (l-C3H) and a cyclic (c-C3H) form, and both
these neutral species have been detected in inter-
stellar molecular clouds.
C2H5OCHO
▶ Ethyl Formate
History
A detection of the linear cation C3H+ (denoted

C3H l-C3H+) in an interstellar ▶ photodissociation
region has been reported by radio astronomers
▶ Propynylidyne (Pety et al. 2012). However, in the absence of
laboratory data on its rotational spectrum, the
identification depended on quantum mechanical
C3H+ calculations and has been challenged. The carrier
of the detected emission lines has subsequently
William M. Irvine been detected in other molecular clouds
University of Massachusetts, Amherst, MA, USA (McGuire et al. 2013). Laboratory spectra and
very accurate theoretical calculations have been
recently reported (Br€unken et al. 2014;
Synonyms Botschwina et al. 2014) confirming C3H+ as the
carrier of the lines observed by radio
Propynylidynium, l-C3H+ astronomers.

DOI 10.1007/978-3-662-44185-5
346 C3H2, c-C3H2
See Also
C4H
▶ Molecular Cloud ▶ Butadiynyl Radical
▶ Photodissociation Region
▶ Propynylidyne
C4H2
References and Further Reading ▶ Diacetylene
Botschwina P, Stein C, Sebald P, Oswald R (2014) Strong

theoretical support for the assignment of B11244 to
l-C3H+. Astrophys J 787:72 C5N
Br€unken S, Kluge L, Stoffels A, Asvany O, Schelemmer
S (2014) Laboratory rotational spectrum of l-C3H+ and
confirmation of its astronomical detection. Astrophys ▶ 4-Cyano-1,3-Butadiynyl
J 783:L4
McGuire BA, Carroll PB, Loomis RA, Blake GA, Hollis
JM, Lovas FJ, Jewell PR, Remijan AJ (2013) A search
for l-C3H+ and l-C3H in Sgr B2(N), Sgr B2(OH), and
the dark cloud TMC-1. Astron Phys J 774: Article 56 C6H6
Pety J, Gratier P, Guzmán V, Roueff E, Gerin M,
Goicoechea JR, Bardeau S, Sievers A, Le Petit F, Le ▶ Benzene
Bourlot J, Belloche A, Talbi D (2012) The IRAM-30 m
line survey of the horsehead PDR. II. First detection of
the l-C3H+ hydrocarbon cation. Astron Astrophys
548:A68
CA
▶ Cyanoacetylene
C3H2, c-C3H2
CAA
▶ Cyclopropenylidene
▶ N-Carbamoyl Amino Acid
CAB, Spain
C3H7CN
Michel Viso
▶ Propyl Cyanide CNES/DSP/SME, Vétérinaire/DVM,
Astro/Exobiology, Paris Cedex 1, France
Definition
C3N
The Spanish Centro de Astrobiologia (CAB) was
▶ Cyanoethynyl Radical created as a Joint Center between CSIC (Consejo
Cahn Ingold Prelog Rules 347
Superior de Investigaciones Cientı́ficas) and the ▶ Extremophiles

Spanish space agency INTA (Instituto Nacional ▶ Genetics
de Técnica Aeroespacial) with the support of the ▶ Geomicrobiology
Comunidad Autónoma de Madrid (CAM). The ▶ Herschel, William
main goal of the CAB is to provide a truly trans- ▶ Mars Science Laboratory
disciplinary research environment for the devel- ▶ NAI
opment of the new science of Astrobiology. The ▶ PLATO 2.0 Satellite C
CAB operates with the specific new contribution of ▶ Terrestrial Analog
a common methodology based on complexity the-
ory and the application of the scientific method to
understanding the origin of life by exploring the
habitability conditions on Earth and beyond, within
the Solar System, or in extrasolar planets. CAB Cahn Ingold Prelog Rules
scientific activities started in late 1999 at temporary
buildings, while awaiting a new building to be Henderson James (Jim) Cleaves II
constructed and equipped. The new CAB premises Earth–Life Science Institute (ELSI), Tokyo
were inaugurated in January 2003. Institute of Technology, Meguro–ku, Tokyo,
CAB is located within the campus of INTA Japan
and, in addition, has one astronomical facility Institute for Advanced Study, Princeton, NJ,
with a robotic telescope at the Observatorio USA
Astronómico Hispano-Alemán de Calar Alto Blue Marble Space Institute of Science,
(CAHA). CAB’s scientists and engineers are Washington, DC, USA
deeply involved in space missions such as MSL, Center for Chemical Evolution, Georgia Institute
▶ ExoMars, INTEGRAL, Bepi Colombo, of Technology, Atlanta, GA, USA
PLATO, HERSCHEL, and SPICA.
Synonyms
History
CIP rules
The origin of the CAB goes back to a proposal
presented to NASA in 1998 by a group of Spanish
and American scientists, led in Spain by Juan
Pérez-Mercader, to join the newly created NASA Definition
Astrobiology Institute (NAI). The group was inte-
grated within the ▶ NAI as a full Associate Mem- In chemistry, these are a set of rules for determin-
ber in 2000. Following an exchange of letters at ing the stereochemistry of a molecule. For mole-
Government level, CAB became the first Associ- cules with double bonds, stereoisomerism can
ate Member of NAI outside the US. The other NAI be E (entgegen, German for opposed) or
Associate Member since 2003 is the Australian Z (zusammen, German for together), also
Centre of Astrobiology. referred to in English as trans and cis, respec-
tively. For stereocenters, molecules are denoted
as S or R (for sinister or rectus) indicating
See Also whether the sequence of groups follows a left or
right direction of viewing, with priority of sub-
▶ Evolution, Biological stituents assigned on the basis of molecular
▶ ExoMars weight.
348 CAIs
See Also irregular, fluffy, round, and compact. Some

CAIs show evidence of melting, probably during
▶ Stereoisomers chondrule formation. Others exhibit evidence of
reprocessing in the chondrite parent bodies (see
entry ▶ Meteorite, Allende). CAIs are enriched in
16O relative to chondrules. Some of them are
enriched in the heavy isotopes of magnesium
CAIs and silicon. They often are isotopically anoma-
lous relatively to the bulk Solar System. These
Matthieu Gounelle anomalies, at the level of a few hundred ppm,
Laboratoire de Minéralogie et Cosmochimie du were found mostly in neutron-rich isotopes, such
Muséum (LMCM) MNHN USM 0205 – CNRS as 48Ca or 50Ti. They contain a high abundance of
UMR 7202, Muséum National d’Histoire short-lived radionuclides (see entry ▶ Cosmo-
Naturelle, Paris, France chemistry). Their Pb-Pb absolute ages
(4,567–4,568 Ma) are older than those of chon-
drules or differentiated meteorites.
Keywords CAIs are very abundant in carbon-rich carbo-
naceous chondrites, while they are virtually
Chondrites; Chronology; Inclusions absent from other chondrites. Studies of CAIs
during the last 30 years are heavily biased toward
CAIs found in the CV3 chondrites epitomized by
Synonyms the Allende meteorite which fell in Mexico in
1969. The reason for that bias is that CAIs in
Calcium-aluminum-rich inclusions CV3 chondrites are large (up to a cm) and abun-
dant (10 % volume). It is, however, important
to note that CAIs in CV3 chondrites are quite
Definition peculiar when compared to CAIs in other chon-
drite groups and may record specific events in the
CAIs (calcium-aluminum-rich inclusions) are solar protoplanetary disk. More studies should
white inclusions found in carbonaceous chon- and will be dedicated to CAIs in other chondrite
drites. They are made of calcium- and groups. Two CAIs were found among the dust
aluminum-rich oxides and silicates. They are brought back by the Stardust spacecraft from
among the older solids of the Solar System. comet Wild 2.
They provide clues on the immediate environ- CAIs probably formed by condensation. Some
ment of the nascent solar system and on its of them endured severe evaporation. Because of
earliest phases. their old age, the extent of the isotopic anomalies
they bear, and their richness in short-lived radio-
nuclides, they are believed to be the first solids to
Overview have formed in the solar protoplanetary disk. As
such, they record the earliest phases of the Solar
CAIs were discovered in the Vigarano ▶ chon- System formation. Short-lived radionuclides as
drite by M. Christophe in 1968. They are an well as isotopic anomalies record nucleosynthe-
assemblage of calcium and aluminum oxides sis processes in generations of stars prior to the
and silicates, i.e., spinel, hibonite, grossite, solar system. Some short-lived radionuclides also
melilite, anorthite, and calcium- and aluminum- record irradiation processes in the early solar
rich pyroxenes. CAIs are enriched in refractory system. Because they formed at high tempera-
elements by a factor of 10–100 relative to bulk ture, in a gas-poor region, they are believed
chondrites. They have variable textures: to have formed close (0.1 AU) to the protosun.
Calvin’s Conception of Origins of Life 349
They were transported to chondrite formation

distances either by winds powered by the mag- Callisto
netic interaction between the Sun and the disk or
by turbulence. Therese Encrenaz
LESIA, Observatoire de Paris - Section de
Meudon, France
See Also C
▶ Cratering Chronology Definition

▶ Cosmochemistry
▶ Meteorite, Allende Callisto is a satellite of Jupiter discovered by
▶ Meteorites ▶ Galileo Galilei in January 1610; it is the outer-
▶ Parent Body most of the Galilean satellites. With a radius of
▶ Protoplanetary Disk 2,410 km, Callisto is, after Ganymede and Titan,
the third biggest satellite in the solar system. Its
distance to ▶ Jupiter is 1,882,700 km or 26 Jovian
radii. Its density is 1.8 g/cm3, typical of icy
Christophe Michel-Lévy M (1968) Un chondre objects. Callisto has been investigated by the
exceptionnel dans la météorite de Vigarano. Bull Soc Voyager 1 and 2 spacecraft in 1979, then by the
Fr Minéral Cristallogr 91:212–214 Galileo orbiter between 1995 and 2003. The sur-
MacPherson GJ, Simon SB et al (2005) Calcium- face of Callisto is heavily cratered and consists of
Aluminium-rich inclusions: major unanswered ques-
tions. In: Krot AN, Scott ERD, Reipurth B (eds) Chon- a mixture of ice and dust. A great basin, Valhalla,
drites and the protoplanetary disk, vol 341. ASP over 500 km in diameter, is the signature of a
Conference Series, San Francisco, pp 225–250 large major impact.
Shu FH, Shang H et al (1996) Toward an astrophysical
theory of chondrites. Science 271:1545–1552
Wadhwa M, Amelin Y et al (2007) From dust to planetes-
imals: implications for the solar protoplanetary disk See Also
from short-lived radionuclides. In: Reipurth VB,
Jewitt D, Keil K (eds) Protostars and planets. Univer- ▶ Galileo Galilei
sity of Arizona Press, Tucson, pp 835–848
▶ Jupiter
Calcareous Sediment
Calvin’s Conception of Origins of Life
▶ Carbonate
Stéphane Tirard
et des Techniques EA 1161, Faculté des
Calcium-Aluminum-Rich Inclusions Sciences et des Techniques de Nantes, Nantes,
France
▶ CAIs
History
Caldophile Melvin Calvin (1911–1997) was an American

biochemist, who discovered, with Andrew Alm
▶ Thermophile Benson (1917–), the cycle of reactions in the
350 Calvin-Benson Cycle
obscure phase of photosynthesis during the 1940s Definition

(Calvin-Benson cycle). Calvin obtained the
Nobel Prize in 1961 for this discovery. A carbon dioxide fixation pathway where a mol-
In 1951, Calvin published one of the first ecule of CO2 condenses with a 5-C compound
works in prebiotic chemistry. He reduced carbon (ribulose 1,5-bisphosphate) to yield two mole-
dioxide in aqueous solution, by ionizing radia- cules of a 3-C compound (3-phosphoglycerate).
tion, to formic acid. These 3-C molecules serve both as precursors for
In 1953, Harold Clayton Urey (1893–1981) biosynthesis and, through a cyclic series of enzy-
rejected this result because of the presence of matic reactions, to regenerate the 5-C molecule
CO2. Urey was in favor of a reductive primitive necessary for the first carboxylating step (Fig. 1).
atmosphere without CO2. However, Calvin The pathway is present in several bacterial line-
maintained his interest for origins of life during ages (e.g., cyanobacteria), and its acquisition by
the rest of his career and published his main book eukaryotic cells (chloroplast in algae and plants)
on this topic in 1969 (Molecular Evolution was through the endosymbiotic association with
towards the Origin of Living Systems on Earth ancient cyanobacteria.
and Elsewhere).
History
See Also Melvin Calvin (1911–1997) and coworkers

established this autotrophic path of carbon in
▶ Calvin-Benson Cycle phototrophic organisms using 14C-labeled carbon
▶ Miller, Stanley dioxide and Chlorella (a green algae) cultures
▶ Urey’s Conception of Origins of Life (Benson and Calvin 1950). Calvin was awarded
with the Nobel Prize in Chemistry in 1961 “for
his research on the carbon dioxide assimilation in
plants.”
Calvin-Benson Cycle
Overview
Juli Peretó
Institut Cavanilles de Biodiversitat i Biologia The Calvin-Benson cycle allows the synthesis of
Evolutiva, Universitat de València, València, one triose from three molecules of carbon dioxide
Spain (Fig. 1): 12 electrons (provided by redox coen-
zymes like NADH or NADPH) and 9 ATP equiv-
alents are required for bringing CO2 to the
Keywords oxidation level of the triose glyceraldehyde
3-phosphate. These fueling requirements are
Biosynthesis; Autotrophy; Carboxylation; Car- supplied by either a phototrophic or a
bon dioxide chemolithotrophic metabolism. The cycle can be
divided into two stages: a reductive carboxylation
of the pentose ribulose 1,5-bisphosphate (RuBP)
Synonyms up to glyceraldehyde 3-phosphate and a series of
rearrangements of carbon skeletons from trioses to
Dark reactions; Reductive pentose phosphate regenerate RuBP throughout 4-, 6-, and 7-C sugar
cycle intermediates (Fig. 1). The first step is catalyzed
Calvin-Benson Cycle 351
3 CO2 6 ATP
3 Ribulose 1,5-BP 6 3-phosphoglycerate

1
2 3 ATP
6 1,3-bisphosphoglycerate
3 Ribulose 5-P
6[H] C
xylulose 5-P + ribose 5-P 6 (Gd3P)
sedoheptulose 7-P Gd3P
DHAP
sedoheptulose 1,7-BP DHAP biomass
xylulose 5-P + erythrose 4-P fructose 1,6-BP
fructose 6-P
Calvin-Benson Cycle, Fig. 1 The Calvin-Benson cycle. chemolithotrophic metabolism. Except for two key enzy-
The stoichiometry of the cycle allows the net synthesis of matic steps (1 and 2), all the transformations are a
one molecule of triose from three molecules of carbon combination of enzymes participating in the Embden-
dioxide (gray boxes). Other five trioses (5 C3) are Meyerhof-Parnas pathway and the non-oxidative
converted into three pentoses (3 C5) necessary to initiate pentosephosphate pathway. Key enzymatic steps:
the cycle again. The energetic cost for the synthesis of one (1) Rubisco; (2) phosphoribulokinase. Abbreviations:
triose is nine ATP molecules and six reducing equivalents Gd3P glyceraldehyde 3-phosphate, DHAP
(NADH or NADPH). These energetic requirements could dihydroxyacetonephosphate
be satisfied by a photosynthetic apparatus or a
by the key carboxylating enzyme ▶ Rubisco. As in cyanobacteria (the group to which the ances-
other carboxylases, Rubisco also shows a prefer- tors of plastids belonged), some aerobic
ence for lighter stable isotopes, and thus CO2 or facultative anaerobic proteobacteria,
fixation results in the depletion of 13C (d13C) in CO-oxidizing mycobacteria, diverse iron-sulfur-
the biosynthesized organic matter ranging from oxidizing firmicutes, and in some green sulfur
20 ‰ to 30 ‰. The second stage of the bacteria. Although Rubisco has been isolated
cycle occurs by the combination of several enzy- from some Archaea, there is no evidence for the
matic activities from the ▶ Embden-Meyerhof- operation of the full cycle in autotrophic species
Parnas pathway and the nonoxidative branch of from this domain (Berg et al. 2010).
the pentose phosphate pathway.
The Calvin-Benson cycle was originally
described in green plant (i.e., chloroplast) photo- See Also
synthesis (Benson and Calvin 1950). It is also
active in endosymbiotic chemolithotrophic ▶ Autotrophy
proteobacteria of invertebrates living in close ▶ Embden-Meyerhof-Parnas Pathway
proximity to hydrothermal vents. In free-living ▶ Isotopic Fractionation (Planetary Process)
prokaryotes, this pathway has been demonstrated ▶ Rubisco
352 Cambrian Explosion
Benson AA, Calvin M (1950) Carbon dioxide fixation by ▶ Burgess Shale Biota
green plants. Annu Rev Plant Physiol 1:25–42
▶ Chengjiang Biota, China
Berg JM, Tymoczko JL, Stryer L (2007) Chapter 20. Bio-
chemistry, 6th edn. Freeman, New York
Autotrophic carbon fixation in archaea. Nat Rev
Cambrian Radiation
Kim BH, Gadd GM (2008) Chapter 10. Bacterial physiol-
ogy and metabolism. Cambridge University Press, ▶ Cambrian Explosion
Cambridge
Campbellrand-Malmani Platform,
South Africa
Cambrian Explosion
Daniele L. Pinti
Thomas H. P. Harvey GEOTOP Research Center for Geochemistry and
Department of Earth Sciences, University of Geodynamics, Université du Québec à Montréal,
Cambridge, Cambridge, UK Montréal, QC, Canada
Synonyms Definition
Cambrian radiation The Campbellrand-Malmani carbonate platform

was deposited on the ▶ Kaapvaal craton, South
Africa, between 2,588 and 2,520 Ma
Definition (Neoarchean). This 1.9-km-thick carbonate plat-
form is preserved over at least 190,000 km2 and
The Cambrian explosion was a major evolution- probably originally covered the entire extent of
ary radiation of organisms, notably multicellular the Kaapvaal craton (~600,000 km2). The
animals (metazoans), that occurred during the Campbellrand-Malmani is the first extensive car-
Cambrian period (542–588 Ma). Precambrian bonate deposit of the Neoarchean prior to the
fossils reveal a mainly microbial biota, with com- buildup of oxygen in the ocean and atmosphere
plex, large-bodied organisms only becoming con- about 2.35 Ga ago. The abundance of aragonite
spicuous during the latest Proterozoic Eon (the pseudomorphs in this unit suggests a neutral to
Ediacaran biota). In contrast, Cambrian fossils slightly alkaline ocean, which in turn may indi-
include abundant evidence for macroscopic cate a low atmospheric CO2 concentration.
motile organisms and unambiguous examples of
most major animal lineages. Exactly what deter-
mined the timing of the Cambrian explosion is
not clear, and current research aims to disentan- See Also
gle the effects of physical environment, ecologi-
cal context, and factors related to development ▶ Calcareous Sediment
and genetics. ▶ Earth’s Atmosphere, Origin and Evolution of
Canadian Precambrian Shield 353
▶ Great Oxygenation Event are surrounded and cut by granites (Hoffman

▶ Ocean, Chemical Evolution of 1989). In detail, the Superior province consists of
▶ Oxygenation of the Earth’s Atmosphere an old (3 Ga) core (NCT) surrounded by linear
belts of volcanic rocks that alternate with granites
and belts of sedimentary rocks which become
younger (2.9 to 2.7 Ga) north and south of the
core (Percival 2007). This architecture suggests the C
Canadian Precambrian Shield Archean provinces (pink on Fig. 1) grew as suc-
cessively younger belts of volcanic and volcano-
Phil Thurston sedimentary rocks added to the continental margin
Laurentian University, Sudbury, ON, Canada by plate-tectonic processes. This was followed by
the intrusion of much granitic rock during the
interval 2.7–2.6 Ga, transforming the Archean
Keywords provinces into a comparatively light, buoyant
▶ craton. The crustal thickness of the cratons is
Accreted terrane; Accretion; Craton; typically 40 km. The cratonization processes
Geological province; Magmatic arc; Superconti- involved the development of large-scale faults
nent cycle that brought gold-bearing fluids from deep in the
Earth to the near-surface environment. Several cra-
tons of this age (2.7 Ga) joined to form the ▶ super-
Synonyms continent “Kenorland” (Aspler and Chiarenzelli
1998). The edges of Kenorland include some
Canadian shield; North American shield much older (3.8 Ga) fragments (red units on
Fig. 1) that were accreted onto the margin of the
supercontinent. Kenorland broke apart by the
development of 2.48–2.10 Ga old rifts containing
Definition dikes and volcanic rocks followed by the deposi-
tion of uranium-bearing, dominantly sandy conti-
The Canadian ▶ Shield is the region of North nental margin sedimentary units [e.g., Hu in brown
America underlain by Precambrian rocks on Fig. 1, on the south edge of the Superior craton].
(>542 Ma), extending from the Arctic Ocean to The various Archean cratons then drifted apart and
the Great Lakes and further south and west in the became separated by ocean basins and island arcs.
subsurface (Fig. 1). The oceans later closed, and the various accreted
terranes and 2.0–1.8 Ga volcanic arcs accreted onto
the margins of the Archean cratons to form the
“Columbia” supercontinent (Zhao et al. 2004).
Overview During the interval 1.8–1.2 Ga, Columbia grew
by the accretion of multiple belts, again consisting
The Canadian Precambrian Shield is subdivided mostly of volcanic and granitic rocks along its
into geological provinces (regions of similar age). southern and eastern margins, e.g., the Yavapai-
The major Archean (>2.5 Ga) provinces are the Central Plains belt [Y on Fig. 1, (1.8–1.7 Ga)], the
Slave (S), Superior (Sup), Rae (R), and Hearne (H). Mazatzal (M) (1.7–1.5 Ga) belt in the USA, the
All of them have a similar architecture: linear Labradorian belt of Quebec and Labrador, and the
sub-provinces or belts of fine-grained sediment 1.3–1.0 Ga Grenville Province (G on Fig. 1)
and subaqueously erupted volcanic rocks which (Karlstrom et al. 1999).
354 Canadian Precambrian Shield
Subdivisions of the Canadian Shield

Archean Cratons and Fragments Paleoproterozoic (2.5-1.6 Ga) Mesoproterozoic (1.6-1.0 Ga)
(1.6-1.0 Ga)
1.8-1.3 Ga Basin 1.3-1.0 Ga Accreted
3.0-2.7 Ga Craton 1.8-0.5 Ga Basin 1.42-1.35 Ga Cover Sequence
−3.9-2.8 Ga Superterrane 1.8 Ga Arc 1.5-1.3 Ga Accreted
−4-3 Ga Basement complex 2.0-1.8 Ga Accreted 1.8-1.7 Ga Accreted
1.95-1.8 Ga Arc
2-1 Ga Arc
2.0-1.8 Ga Arc/oceanic
2.1-1.9 Ga Arc/oceanic
2.4-2.0 Ga Accreted terrane
2.4-2 Ga Accreted
2.4-2.0 Ga Platform/rift
Canadian Precambrian Shield, Fig. 1 Subdivisions of S Slave craton, R Rae craton, H Hearn craton, Wy Wyo-
the Canadian Shield of North America based on the age of ming craton, GC Grouse Creek block. Paleoproterozoic
the rock unit. Archean cratons and subdivisions within: (2.5–1.6 Ga) tectonic domains after Ross (2002) and
Sup Superior craton, NSS Northern Superior superterrane Aspler et al. (2003): Hu Huronian, T Taltson, B (brown)
and Hudson Bay terrane, RA Riviere Arnaud terrane, NCT Buffalo Head, B (green) Great Bear, H Hottah, Fs Fort
North Caribou terrane, M Marmion terrane, N Nain craton, Simpson, W Wabamun, Chinchaga and Ksituan terranes
Carbamonitrile 355
See Also
Cap Carbonates
▶ Archean Eon
▶ Craton Emmanuelle J. Javaux
▶ Proterozoic Eon Palaeobiogeology-Palaeobotany-
▶ Shield Palaeopalynology, Geology Department,
▶ Supercontinent Université de Liège, Liège, Belgium C
References and Further Reading Definition
Aspler LB, Chiarenzelli JR (1998) Two Neoarchean ▶ Cap carbonates are a special type of laminated
supercontinents? Evidence from the carbonate rocks (limestone and/or dolostone)
paleoproterozoic. Sediment Geol 120:75–104
associated with glacial deposits in the
Aspler LB, Pilkington M, Miles WF (2003) Interpretations
of Precambrian basement based on recent aeromag- Neoproterozoic (▶ Snowball Earth). They
netic data, Mackenzie Valley, Northwest Territories, sharply overly (“cap”) unsorted glacial sediments
Geological Survey of Canada, Current Research 2003- called tillites. Their origin is not well understood,
C2, Ottawa, p 11
but they may be related to a particular ocean and
Hoffman PF (1989) Precambrian geology and tectonic
history of North America. In: Bally AW, Palmer AR atmosphere chemistry with high CO2 concentra-
(eds) The geology of North America; an overview. tion and alkalinity from silicate weathering or to
A. Decade of North American geology. Geological upwelling of highly alkaline deep waters follow-
Society of America, Boulder, pp 447–512
ing deglaciation. They are associated with nega-
Karlstrom KE, Harlan SS, Williams ML, McLelland J,
Geissman JW, Ahall K-I (1999) Refining Rodinia; tive incursions of carbon isotopes that are
geologic evidence for the Australia-Western U.S. widespread and can serve for correlation.
connection in the Proterozoic. GSA Today 9:1–7
Percival JA (2007) Geology and metallogeny of the supe-
rior province, Canada. In: Goodfellow W (ed) Mineral
deposits of Canada: a synthesis of major deposit types, See Also
District metallogeny, the evolution of geological prov-
inces and exploration methods, Special publication no 5, ▶ Glaciation
Geological Association of Canada Mineral Deposits
▶ Sedimentary Rock
Division, p 903–928
Ross GM (2002) Evolution of precambrian continental ▶ Snowball Earth
lithosphere in western Canada; results from lithoprobe
studies in Alberta and beyond. Can J Earth Sci
39:413–437
Zhao G, Sun M, LI S, Wilde SA (2004) A Paleo-
Mesoproterozoic supercontinent: assembly, growth Capsid Encoding Organism
and breakup. Earth Sci Rev 67:91–123
▶ Virus
Canadian Shield Carbamide
▶ Canadian Precambrian Shield ▶ Urea
Canyon Carbamonitrile
▶ Chasma, Chasmata ▶ Cyanamide

356 Carbene
Carbene Carbodiimide
Steven B. Charnley William M. Irvine

Solar System Exploration Division, Code 691, University of Massachusetts, Amherst, MA, USA
Astrochemistry Laboratory, NASA Goddard
Synonyms
HNCNH
Definition
A carbene is an organic molecule having two

electrons available on a carbon atom for chemical Definition
bonding. Short-chain and long-chain carbenes are
very common in interstellar chemistry, e.g., C3, Carbodiimide, HNCNH, is a higher-energy iso-
H2CCC, and H2CCCC (Cernicharo et al. 1991a, mer of ▶ cyanamide, H2NCN. The latter is pre-
b, 2000), and even longer chains such as H2C5 sent in interstellar “hot cores,” regions in
and H2C6 (McCarthy et al. 1997). molecular clouds where massive young stars are
forming, and is an effective condensation agent in
prebiotic chemistry (Nooner et al. 1973).
See Also
History
▶ Interstellar Chemical Processes
▶ Interstellar Medium Carbodiimide was detected by radio astronomers
▶ Molecules in Space (McGuire et al. 2012) in the Galactic Center
molecular cloud Sagittarius B2(N).
References
See Also
Cernicharo J, Gottlieb CA, Guélin M et al (1991a) Astro-
nomical detection of H2CCC. Astrophys J 368:L39
Cernicharo J, Gottlieb CA, Guélin M et al (1991b) Astro-
▶ Cyanamide
nomical detection of H2CCCC. Astrophys J 368:L43 ▶ Hot Core
Cernicharo J, Goicoechea JR, Caux E (2000) Far-infrared ▶ Molecules in Space
detection of C3 in Sagittarius B2 and IRC +10216.
Astrophys J 534:L199
McCarthy MC, Travers MJ, Kovacs A (1997) Detection
and characterization of the cumulene carbenes H2C5
and H2C6. Science 275:518
McGuire BA, Loomis RA, Charness CM, Corby JF, Blake
GA, Hollis JM, Lovas FJ, Jewell PR, Remijan AJ
(2012) Interstellar carbodiimide (HNCNH): a new
astronomical detection from the GBT PRIMOS survey
via maser emission features. Astrophysics J758:
L33–L37
Carbimide Nooner DW, Sherwood E, More MA, Oro J (1973) Cyan-
amide mediated syntheses under plausible primitive
▶ Cyanamide earth conditions. J Mol Evol 10:211–220
Carbohydrate 357
carbohydrates that contain an aldehyde group.

Carbohydrate In the structures below, each line represents a
covalent bond and each intersection of lines rep-
Heshan Grasshopper Illangkoon resents an unwritten carbon (C).
Department of Chemistry, University of Florida, There are 2n stereoisomers of a given sugar,
Gainesville, FL, USA where n is the number of ▶ chiral centers in the
molecule. For instance, the aldopentoses contain C
five carbons, three of which are chiral centers,
Keywords giving 23 = 8 stereoisomers. These pentoses
have the common names of ribose, arabinose,
Cellulose; Formaldehyde; Formose; Fructose; xylose, and lyxose and each come in a d or l
Glucose; Glyceraldehyde; Glycolaldehyde; Ori- ▶ enantiomer. The d or l configuration of the
gins of life; Primordial soup; Ribose; Sucrose; sugar is determined by the position of the alcohol
Sugars; Disaccharide; Monosaccharide; Oligo- on the chiral center furthest away from the alde-
saccharide; Polysaccharide hyde carbon. If the alcohol is on the right of the
backbone, the sugar is classified as d, while if it is
on the left, it is l. In the case of aldopentoses, this
Synonyms chiral center is on carbon four (Fig. 2). On Earth,
the d isomer prevails as the dominant carbohy-
Saccharide drate incorporated in biological systems; how-
ever, it has been postulated that l sugars could
also be used as the predominant isomer in alter-
Definition nate “alien” biochemistries.
Carbohydrates can also form cyclic structures
Carbohydrates are molecules composed solely of that produce an anomeric center giving the sugar an
carbon, oxygen, and hydrogen with the general alpha (alcohol pointing down) or beta (alcohol
formula Cn(H2O)n. The ratio of carbon to hydro- pointing up) label. Ribose, for example, can be
gen to oxygen atoms in a carbohydrate is 1:2:1. found in a furanose (five-membered) or pyranose
Carbohydrates are involved in cell signaling, (six-membered) form. Both the alpha and beta
serve as a source of energy, and provide structure configurations of the furanose and pyranose forms
to cells. Chains that alternate negatively charged of ribose are shown in Fig. 3. The ribose found in
phosphates with carbohydrates form the back- DNA is in a furanose configuration, whereas ribose
bones of the genetic biopolymers RNA and DNA. in aqueous solutions and crystalline form is pre-
dominantly pyranose (Šišak et al. 2010).
Overview Presence of Carbohydrates in the Universe

Astrobiologists studying the ▶ origins of life seek
Carbohydrates are an essential component of life an answer to fundamental questions related to the
on Earth. From ▶ ribose being an integral com- origins of carbohydrates. Through the use of
ponent of the genetic biopolymers DNA and radio telescopes, scientists have determined the
RNA to polymers of glucose used for cell wall presence of carbohydrates and their building
support in plants, carbohydrates are involved in blocks in interstellar gas clouds. Examples of
many crucial biological functions. The complex some organic molecules reported in these
structures of carbohydrates can be easily visual- clouds (Fig. 4) include ▶ formaldehyde,
ized using ▶ Fischer projections. Figure 1 depicts ▶ glycolaldehyde, and dihydroxyacetone
Fischer projections of aldoses, a class of (Snyder et al. 1969; Hollis et al. 2004; Weaver
358 Carbohydrate
CHO CHO
H OH HO H
H OH H OH
CH2OH CH2OH
D-erythrose D-threose
CHO CHO CHO CHO
H OH HO H H OH HO H
H OH H OH HO H HO H
H OH H OH H OH H OH
CH2OH CH2OH CH2OH CH2OH

D-ribose D-arabinose D-xylose D-lyxose
CHO CHO CHO CHO CHO CHO CHO CHO
H OH HO H H OH HO H H OH HO H H OH HO H
H OH H OH HO H HO H H OH H OH HO H HO H
H OH H OH H OH H OH HO H HO H HO H HO H
H OH H OH H OH H OH H OH H OH H OH H OH
CH2OH CH2OH CH2OH CH2OH CH2OH CH2OH CH2OH CH2OH
D-allose D-altrose D-glucose D-mannose D-gulose D-idose D-galactose D-talose
Carbohydrate, Fig. 1 The four-, five-, and six-carbon d-aldoses
Carbohydrate, 1 1 1
CHO CHO CH2OH
Fig. 2 Carbohydrates can
occur in either D or 2 2 2
H OH H OH O
L forms. The ketose
ribulose has an oxidized 3 3 3
center at carbon two H OH H OH H OH
4 4 4
HO H H OH H OH
5 5 5
CH2OH CH2OH CH2OH
L-ribose D-ribose D-ribulose

aldose aldose ketose
and Blake 2005; although the detection of dihy- Prebiotic Synthesis of Carbohydrates
droxyacetone has been questioned – see ▶ Mole- The prebiotic synthesis of carbohydrates is fun-
cules in Space). Meteoritic bombardment could damental to astrobiology and origins of life
have delivered these molecules to the surface research. The “▶ RNA World” hypothesis,
of the early Earth (Sephton 2002). Aldol reac- which purports that life evolved from simple
tions of glycolaldehyde and formaldehyde on a RNA molecules, presupposes the existence of a
primitive Earth could also have been sources of prebiotic soup with abundant building blocks,
glyceraldehyde, erythrulose, and higher carbohy- including amino acids, carbohydrates, purines,
drates (Fig. 5). and pyrimidines.
Carbohydrate 359
Carbohydrate, HO HO
Fig. 3 The furanose (five- OH β
membered) and pyranose O O
(six-membered) forms of
d-ribose and their alpha (a)
OH α
and beta (b) anomers
OH OH OH OH
furanose C
O O OH β
OH OH α OH
OH OH OH OH
pyranose
Carbohydrate, CHO
O O
Fig. 4 Formaldehyde and O H
some lower carbohydrates C H OH
glycolaldehyde,
H H HOH2C CH2OH
d-glyceraldehyde, and CH2OH CH2OH
dihydroxyacetone
formaldehyde glycolaldehyde D-glyceraldehyde dihydroxyacetone
−
O O O O
H
H
H H H H H H OH
OH OH formaldehyde OH
glycolaldehyde ene-diol of glycolaldehyde glyceraldehyde
Carbohydrate, Fig. 5 An example of an aldol addition of formaldehyde to glycolaldehyde
One of the first reported instances of prebiotic One argument against the RNA world arising
carbohydrate synthesis occurred in 1861 when a from the ▶ primordial soup involves the forma-
German scientist, Butlerov, added formaldehyde tion of “problematic tar,” which is a seemingly
to hot solutions of barium and calcium useless by-product of highly reactive species
hydroxide forming a sweet sugary substance (Larralde et al. 1995). The propensity of the
(Butlerov 1861). This became known as the formose reaction to form tar is an indication of
▶ formose reaction. A century later, Breslow dis- the functionality and reactivity of these building
covered that the glycolaldehyde formed in this blocks. The tar prevents the accumulation of
reaction initiates a series of autocatalytic reaction genetically relevant carbohydrates and compli-
cycles which, over time, fix more formaldehyde cates in-depth product analysis. Recently, how-
to give higher carbohydrates (Breslow 1959). ever, ▶ borate minerals have been shown to
A side reaction in this experiment is the dispro- stabilize and direct the reactivity of carbohy-
portionation of ▶ formaldehyde to form ▶ meth- drates, making them potentially important partic-
anol and formic acid in a process now known as ipants in prebiotic reactions (Ricardo et al. 2004).
the Cannizzaro reaction (Fig. 6). Here, borate binds to adjacent alcohols of sugars
360 Carbon
− H
O O O O
− strong base −
OH H H + H O
H H H H H O −
O H
−
formaldehyde formaldehyde formic acid methanol
Carbohydrate, Fig. 6 In the Cannizzaro reaction, formaldehyde reacts in strong base to give formic acid and methanol
(both shown in their deprotonated forms)
as they are formed. This in turn reduces their Hollis JM et al (2004) Green bank telescope observations
reactivity and improves carbohydrate stability, of interstellar glycolaldehyde: low temperature sugar.
Astrophys J 613:L45–L48
preventing further uncontrolled tar-forming Larralde R, Robertson MP, Miller S (1995) Rates of
reactions. decomposition of ribose and other sugars: implications
In a different approach to prebiotic carbohy- for chemical evolution. Proc Natl Acad Sci U S A
drate synthesis, activated pyrimidine nucleosides 92:8158–8160
Powner MW, Gerland B, Sutherland JD (2009) Synthesis of
are assembled, sidestepping the model of adding activated pyrimidine ribonucleotides in prebiotically
a nucleobase to a previously synthesized ribose plausible conditions. Nature 459:239–242
sugar (Powner et al. 2009). Though this method- Ricardo A, Carrigan M, Olcott AN, Benner SA
ology does not yet account for the synthesis of (2004) Borate minerals stabilize ribose. Science
303:196
purine nucleosides, it opens new avenues for Sephton M (2002) Organic compounds in carbonaceous
research toward the prebiotic synthesis of RNA meteorites. Nat Prod Rep 19:292–311
and DNA. Šišak D, McCusker LB, Zandomeneghi G, Meier BH,
Bläser D, Boese R, Schweizer WB, Gilmour R, Dunitz
JD (2010) The crystal structure of D-ribose – at last!
Angew Chem Int Ed 49:4503–4505
See Also Snyder LE, Buhl D, Zuckerman B, Palmer P (1969)
Microwave detection of interstellar formaldehyde.
▶ Aldose Phys Rev Lett 22:679
Weaver SLW, Blake GA (2005) 1, 3-Dihydroxyacetone in
▶ Biopolymer Sagittarius B2(N-LMH): the first interstellar ketose.
▶ Borate Astrophys J 624:L33
▶ Disproportionation
▶ Fischer Projection
▶ Formaldehyde
▶ Formose Reaction Carbon
▶ Glycolaldehyde
▶ Ketose Alan W. Schwartz
▶ Methanol Radboud University Nijmegen, Nijmegen,
▶ Origin of Life The Netherlands
▶ Primordial Soup
▶ Ribose
▶ RNA World Synonyms
Elemental carbon
Breslow R (1959) On the mechanism of the formose

reaction. Tetrahedron Lett 1:22–26
Definition
Butlerov A (1861) Formation synthétique d’une substance
sucrée. C R Acad Sci 53:145–147 Carbon: The sixth element of the periodic table
Carbon 361
Overview carbon, 12C. While the isotopic fractionation

ratio (d13C) was once thought to be a reliable
Carbon is the sixth element in the periodic table. indication of a biological carbon fixation
Its stable isotopes include 12C, the nucleus of (Schidlowski et al. 1983), this has recently
which contains six protons and six neutrons, and been questioned (McCollom and Seewald
13
C, which contains six protons and seven neu- 2006).
trons. Carbon is the fourth most abundant ele- C
ment in the solar atmosphere, after hydrogen, Electronic Structure
helium, and oxygen. This predominance is The electronic structure of carbon is abbreviated
largely due to the autocatalytic role of carbon in as 1s2, 2s2, 2p2 (two electrons in the 1s orbital,
the stellar nucleosynthesis of elements higher two electrons in the 2s orbital, and two electrons
than hydrogen via proton addition (Bethe 1939): in the 2p orbital). The element can therefore
accept four electrons in the outermost (valance)
12
C þ H!13 N þ g shell to complete an octet and form a neon-like
configuration: 1s2, 2s2, 2p6. It was pointed out by
13
N!13 C þ eþ
Wald that the prevalence of H, O, N, and C in
13
C þ H!14 N þ g biochemistry can be rationalized by the fact that
14
N þ H!15 O þ g these elements are the smallest that can reach
15
O!15 N þ eþ stable (filled, i.e., noble gas-like) electronic con-
figurations in the valance shell by adding (i.e.,
15
N þ H!12 Cþ4 He sharing with other atoms) one, two, three, and
four electrons, respectively, and thereby forming
Carbon Isotopes covalent bonds (Wald 1958).
Carbon exists on Earth in the form of two stable This suggests that the same properties of these
(12C and 13C) and one unstable isotope (14C). elements would similarly be responsible for a
12
C and 13C have abundances of 98.89 % and central role for organic chemistry (and biochem-
1.108 %, respectively (Wederpohl 1978). The istry) anywhere in the universe. Wald also argued
standard atomic weight of carbon on Earth that silicon’s electronic configuration (one period
(abridged to five figures) is, therefore, 12.011 lower in the periodic table; 1s2, 2s2, 2p6, 3s2, 3p2),
(Commission on Isotopic Abundances and while potentially also permitting the forming of
Atomic Weights, IUPAC). 14C (six protons and covalent bonds by accepting four electrons in the
eight neutrons) is produced in the upper atmo- outermost shell, is substantially larger than car-
sphere by the interaction of thermal neutrons bon. The larger distance between the nucleus of
(produced from cosmic rays) and nitrogen silicon and the outer electronic orbitals results in
atoms. The isotope has a half-life for decay differences in bonding energies, as well as the
(into 14N and a beta particle, b) of 5,730 years geometry of the orbitals, compared to carbon.
and is employed in determining the age of The result is that C forms stable covalent bonds
objects having a biological origin up to about (single as well as multiple) with itself and with
50,000 years of age (Arnold and Libby 1949). In other atoms, while Si has much less tendency to
addition to the use of 14C decay kinetics to do so. Thus, CO2 is a discrete molecule and a
determine the age of objects of archeological stable gas, and C-C bonds produce stable chains
and anthropological interest, the ratio of the of hydrocarbons. SiO2, on the other hand, is a
stable isotopes 12C/13C is an important indica- crystalline solid (of average composition SiO2),
tion of the pathway of synthesis of organic com- which exists as an unlimited three-dimensional
pounds. Because of the kinetic isotope effect, network, and Si-Si chains are not stable in the
both biological and nonbiological synthetic pro- presence of water or O2. Carbon is thus regarded
cesses can produce organic material containing by most astrobiologists as being central to and
a distinctive enrichment of the lighter isotope of probably necessary for life.
362 Carbon
Native Forms of Carbon Compounds of Carbon and the History of

In spite of its importance to life and its role in Organic Chemistry
the biosphere, elemental carbon is only a trace Although the element has been known from
constituent in most of the Earth’s crust, where ancient times, an understanding of its chemical
it is present at an average level of 200 ppm. nature and particularly its role in organic material
In sedimentary rocks, carbon is abundant but did not emerge until the development of analyti-
usually in compounds such as carbonates cal methods of analysis by Lavoisier and others in
(limestone) or organic matter (petroleum, the eighteenth century. Thus, Lavoisier demon-
coal). In the biosphere, hydrosphere, and atmo- strated that carbonic acid or carbon dioxide was
sphere, carbonic acid equilibria (carbon dioxide, formed by the union of carbon and oxygen or by
carbonate, and bicarbonate) play major roles in burning organic substances in air or oxygen
the weathering of crustal rocks and presumably (Moore 1918). The terms “organic chemistry”
would have similar roles on other, Earth-like and “organic compound” were historically
planets. intended to indicate the nature of biologically
The primary forms of elemental carbon in derived compounds of carbon, which were
Earth’s crust and mantle are graphite and dia- thought to be unique to life. Today, organic
mond. Diamond is thought to form under high- chemistry is usually described as the chemistry
pressure and high-temperature conditions in the of compounds of carbon but probably can be
mantle and to be transported to the surface by more accurately regarded as focusing on the
magmatic activity. Graphite has a sheet-like net- chemistry of hydrocarbons (compounds com-
work of so-called sp2 (planar)-bonded carbon posed of hydrogen and carbon) and of their deriv-
atoms (see below) and can be hydrogenated to atives with oxygen and nitrogen.
produce hydrocarbons or oxidized to carboxylic Modern organic chemistry is generally consid-
acids. In contrast, diamond, which has a tetrahe- ered to have started in the mid-nineteenth century
dral or sp3 structure, forms face-centered cubic with the work of Wöhler, who synthesized the
crystals which are difficult to oxidize or reduce. well-known animal waste product, urea. The
The hexagonal mineral lonsdaleite, another allo- actual story of Wöhler’s work is somewhat
trope which is said to be even harder than dia- more complicated than the usual description.
mond, is only found in strongly shocked areas The great chemist was attempting to synthesize
such as meteorite craters, where it is thought to ammonium cyanate by studying the reaction
have formed from graphite during impact between cyanogen and (aqueous) ammonia, one
(Wederpohl 1978). Similarly, fullerenes and of the products of which was a white crystalline
related “cage” forms of carbon also require substance. The same product was also obtained
exceptional conditions to form but have been by the reaction of lead cyanate and ammonia. It
identified in the unshocked Allende and Murchi- was in the course of purifying the latter that
son meteorites as C60–C400 fullerenes Wöhler obtained urea. His realization of the
(Becker et al. 2000). Graphene, a linear sheet potential importance of the reaction led him to
structure consisting of hexagonal lattices of car- write to Berzelius, saying that he could make urea
bon has long been recognized as a theoretical without using a kidney or even an animal (McKie
possibility, but was only synthesized in 2004 1944). In spite of the enthusiastic nature of this
(Novoselov et al. 2004). It forms the basis for a communication to Berzelius, the latter’s reaction,
new series of sheet structures and can be thought as well as those of many of Wöhler’s contempo-
of as the basic unit of fullerenes. Similarly, car- raries, was restrained and ▶ vitalism only gradu-
bon nanotubes have been described in which ally died out.
graphene sheets comprise the basic structural During this same period, chemists became
unit. For a review of these structures see Hirsch aware of the existence of meteorites containing
(Hirsch 2010). organic compounds. Berzelius and Wöhler,
Carbon 363
among others, remarked on the presence of asymmetric center. More complex molecules,
organic material in several carbonaceous meteor- such as the biological product tartaric acid,
ites. It is noteworthy, however, that neither Ber- which led Pasteur to his discovery, contain more
zelius nor Wöhler thought that the organic than one chiral center and can produce even more
compounds present were necessarily biological complicated sets of chiral enantiomers.
products (Nagy 1975). Today, the universality
of organic chemistry has become obvious, not C
only through the investigation of meteorites and See Also
comets but due to observations of interstellar
organic compounds. ▶ Carbonaceous Chondrite
▶ Carbon Isotopes as a Geochemical Tracer
Molecular Structure of Carbon Compounds ▶ Chirality
Bond formation between carbon atoms (or with ▶ Comet
other elements) occurs by sharing electrons and ▶ Delta, Isotopic
involves a process known as hybridization of ▶ Fischer-Tropsch Effects on Isotopic
molecular orbitals. In methane and related hydro- Fractionation
carbons, the carbon atom is located in the center ▶ Interstellar Chemical Processes
of a tetrahedron, with the bonded hydrogen atoms ▶ Organic Molecule
located at the corners. This geometry was ▶ Vitalism
explained by Pauling:
We have thus derived the result that an atom in
which only s and p eigenfunctions contribute to References and Further Reading
bond formation and in which the quantization in
polar coordinates is broken can form one, two, Arnold JR, Libby WF (1949) Age determinations by
three, or four equivalent bonds, which are directed radiocarbon content: checks with samples of known
toward the corners of a regular tetrahedron. age. Science 110:678–680
(Pauling 1931) Becker L, Poreda RJ, Bunch TE (2000) Fullerenes: an
extraterrestrial carbon carrier phase for noble gases.
The tetrahedral geometry of compounds of Proc Natl Acad Sci 97:2979–2983
Bethe HA (1939) Energy production in stars. Phys Rev
carbon thus results from so-called sp3 hybridized
55:434–456
orbitals. As a consequence of this symmetry, Henderson LJ (1913) The fitness of the environment: an
substitution of other atoms bonded to the central inquiry into the biological significance of the proper-
carbon atom (such as substitution of the hydrogen ties of matter. Macmillan, Boston
Hirsch A (2010) The era of carbon allotropes. Nature
atoms in methane, CH4 by atoms or groups A, B,
Materials 9:868–871
and C to produce CHABC), produces an unsym- McCollom TM, Seewald JS (2006) Carbon isotope com-
metrical molecule which displays the phenome- position of organic compounds produced by abiotic
non of ▶ chirality. Synthesis of such a chiral synthesis under hydrothermal conditions. Earth Planet
Sci Lett 243:74–84
molecule by a spontaneous chemical reaction
McKie D (1944) Wöhler’s ‘synthetic’ urea and the rejec-
produces a mixture of isomers or enantiomers tion of vitalism: a chemical legend. Nature
which are chemically but not physically identical. 153:608–610
It can readily be seen by inspecting models that Moore FJ (1918) A history of chemistry. McGraw-Hill
Books, New York, Reprint by General Books, Mem-
such simple examples of chiral enantiomers are
phis, Tennessee, 2009
mirror images of each other. The phenomenon of Nagy B (1975) Carbonaceous meteorites. Elsevier,
chirality was discovered historically by Pasteur, Amsterdam, pp 43–78
in the form of “optical activity,” as determined by Novoselov et al. (2004) Electric Field Effect in Atomi-
cally Thin Carbon Films. Science 306:666–669
the property of rotating the plane of polarization Pauling L (1931) The nature of the chemical bond. Appli-
of a plane-polarized beam of light. The carbon cations of results obtained from the quantum mechan-
atom in this simple compound is referred to as an ics and from a theory of paramagnetic susceptibility to
364 Carbon Concentrating Mechanisms
the structure of molecules. J Am Chem Soc Overview

53:1367–1400
Schidlowski M, Hayes JM, Kaplan IR (1983) In: Schopf
JW (ed) Isotopic inferences of ancient biochemistries: Carbon is the fourth commonest element in the
carbon, sulfur, hydrogen, and nitrogen in earth’s earli- Universe and is present in all five Earth spheres
est biosphere. Princeton University Press, Princeton, (▶ biosphere, geosphere, pedosphere, ▶ hydro-
pp 149–186 sphere, and atmosphere). The annual rates of bio-
Wald G (1958) Introduction to the 1958 edition of Hen-
derson. Beacon Press, Boston, pp xvi–xxiv, op.cit logical carbon cycling are several orders of
Wederpohl KH (ed) (1978) Handbook of geochemistry, magnitude greater than geological cycling of car-
vol II/1(6). Springer, Berlin bon and very responsive to environmental fluctu-
ations. Carbon is essential to life on Earth, and the
biological processes of light-mediated carbon fix-
ation (photosynthesis, chemolithoautotrophy),
Carbon Concentrating Mechanisms ▶ aerobic respiration of organic carbon to form
inorganic CO2, and (to a lesser extent) ▶ anaerobic
▶ Carboxysomes, Structure and Function respiration to form methane (and some CO2) are
important mechanisms for processing carbon
within the five spheres. Exchange of carbon
between land and atmosphere is driven by the
Carbon Cycle, Biological key biological processes, but in the oceans is dom-
inated by physical exchange as the world’s oceans
J. Cynan Ellis-Evans hold vast quantities of dissolved inorganic carbon.
UK Arctic Office, Strategic Coordination Group, This reservoir of dissolved inorganic carbon (DIC)
British Antarctic Survey, Cambridge, UK is relatively dynamic, but other reservoirs are more
stable. In the case of terrestrial carbon cycling, low
rates of decomposition can result in accumulation
Keywords of organic carbon (e.g., peat deposits, bogs) which
can under certain geological conditions form sub-
Biogeochemical cycles; Carbon source; Fermen- stantial coal, oil, and gas (including methane
tation; Photosynthesis; Respiration hydrate) deposits. In the oceans, sedimentation of
plankton with calcareous skeletons results in
burial and limestone formation though this can
Definition also form through reaction of carbonate ions with
calcium. This range of organic and particularly
The carbon cycle is one of the most important inorganic carbon deposits represents significant
biogeochemical cycles on Earth and involves all long-term sinks for atmospheric CO2. Anthropo-
five environmental spheres. Its geological com- genic activities significantly contributing to car-
ponents operate on a scale of millions of years, bon cycling include burning of fossil fuels and
while biological carbon cycling operates over a agro fuels and industrial processes, such as cement
scale of days to thousands of years. Every organ- production (limestone decomposition). The
ism on Earth needs carbon either for structure or observed buildup of CO2 in the atmosphere since
energy, and the primary biological processes in the start of the industrial revolution and associated
carbon cycling are ▶ photosynthesis and respira- regional warming has led to warmer ocean surface
tion. Incorporation of biological carbon in sedi- waters. Warmer waters absorb less CO2, and so the
mentary deposits subsequently used as fossil “braking” effect of the oceans on rising atmo-
fuels is now also an important process in biolog- spheric CO2 concentrations is reduced. Warming
ical carbon cycling, notably through influencing also allows potential release of methane from
the global carbon budget and thus the Earth’s temperature-sensitive methane hydrate deposits
climate. within shallow marine sediments and from
Carbon Dioxide 365
thawing permafrost peat deposits on land, as well symmetrical and nonpolar. The triple point of
as extending the active season for soil respiration, pure CO2 occurs at 56.6 C and 518 kPa
further enhancing release of carbon dioxide and (Fig. 1). As a gas, carbon dioxide is clear, color-
methane to the atmosphere. less, and odorless.
See Also C
Overview
▶ Aerobic Respiration
Carbon dioxide occurs predominantly as either a
gas or a solid ice in the solar system. As a gas, it is
▶ Biosphere
the predominant constituent of the atmospheres
▶ Fermentation
of Venus and Mars, as well as a minor component
▶ Hydrosphere
of the atmospheres of many other planetary bod-
▶ Methanogens
ies including the Earth. As an ice, CO2 is a sig-
▶ Photosynthesis
nificant component of comets and the Martian
polar caps. Liquid carbon dioxide requires tem-
References and Further Reading peratures above 56 C and pressures greater
than 5 bar (Fig. 1) to exist. Such conditions are
Knonova MM (1966) Soil organic matter. Pergamon, New not presently known to occur in the solar system,
York
but might have been present on early Mars and
Olson JS, Pfuderer HA, Chan Y-H (1978) Changes in the
global carbon cycle and the biosphere, ORNL/EIS- may occur on extrasolar planets. Carbon dioxide
109. Oak Ridge National Laboratory, Tennessee has been found in interstellar clouds in gas and
Sedjo R (1993) The carbon cycle and global forest eco- ice phases by the Infrared Space Observatory
system. Water Air Soil Pollut 70:295–307
(ISO).
Carbon dioxide is also present in the interior of
the Earth, where it occurs as a trace component
Carbon Dioxide within the crystal structures of minerals or
occupies void spaces between minerals. At the
Thomas McCollom oxidation state of the Earth’s mantle, carbon
Laboratory for Atmospheric and Space Physics, dioxide together with the minerals graphite and
University of Colorado, Boulder, CO, USA diamond is the predominant stable form of car-
bon. Carbon dioxide dissolves into magmas as
they form within the Earth’s interior and is
Keywords released into the atmosphere when the magmas
cool and solidify as volcanic rocks at the surface.
Carbon cycle; Carbon fixation; Carbon-silicate As a consequence, carbon dioxide is a major
cycle; Greenhouse effect component of volcanic gases and is the primary
form of carbon in those gases. Carbon dioxide
that accumulates in the atmosphere contributes to
Synonyms the ▶ greenhouse effect that plays a significant
role in warming of planetary surfaces.
CO2 Carbon dioxide dissolves readily in water,
where it hydrates to form carbonic acid
(H2CO3(aq)), which in turn generates acidity
Definition through the release of protons (H2CO3(aq) ! H+
+ HCO3 ! 2H+ + CO32). The protons released
Carbon dioxide is a triatomic compound with the can react with minerals to cause rock weathering,
chemical composition CO2. The molecule is altering the rock’s mineralogy and releasing
366 Carbon Isotopes as a Geochemical Tracer
Carbon Dioxide, 10,000

Fig. 1 Phase diagram for uid
+ liq
pure carbon dioxide Solid
1,000 CO2 solid
CO2 liquid
100
Pressure (atm)
Sublimation point
–78.5°C at 1 atm + gas
10 Liquid
Critical point
1
Triple point
0.1
s
–56.6°C at 5.11 atm
ga
+
lid
So
0.01 CO2 gas
0.001
−140 −120 −100 −80 −60 −40 −20 0 20 40 60 80 100
Temperature (°C)
Pressure-temperature phase diagram for CO2
cations such as Ca2+ and Mg2+ into solution. ▶ Photoautotroph

Under some circumstances, the cations released ▶ Reduction
can react with dissolved CO2 to precipitate as ▶ Respiration
carbonate minerals including calcite (CaCO3)
and siderite (FeCO3) as part of the carbon-silicate
cycle. The net result is the removal of atmo- References and Further Reading
spheric CO2 into carbonate-bearing rock.
Some biological organisms, the photoauto- Archer D (2010) The global carbon cycle. Princeton Uni-
versity Press, Princeton
trophs and chemoautotrophs, are capable of
MacKenzie FT, Lerman A (2006) Carbon in the
transferring electrons to carbon dioxide (CO2 geobiosphere – earth’s outer shell. Springer, Dordrecht
▶ reduction) to form bioorganic compounds, a
process termed carbon fixation. Conversely,
heterotropic organisms consume bioorganic mat-
ter and convert much of it back to carbon dioxide
in a process known as ▶ respiration. Transforma- Carbon Isotopes as a Geochemical
tion of CO2 through carbon fixation and respira- Tracer
tion is a primary component of the biological
carbon cycle. Noah Planavsky1, Camille Partin2 and
Andrey Bekker1
1
Department of Geology and Geophysics, Yale
See Also University, New Haven, CT, USA
2
Department of Geological Sciences, University
▶ Carbon Cycle, Biological of Manitoba, Winnipeg, MB, Canada
▶ Carbon Monoxide
▶ Carbonate
▶ Chemoautotroph Keywords
▶ Infrared Space Observatory Carbon cycle; Carbonates; Organic carbon burial
Carbon Isotopes as a Geochemical Tracer 367
Definition on Earth possess d13C values ranging from

35 ‰ to +5 ‰, which reflect the typical kinetic
Carbon has two stable isotopes (12C and 13C) and isotope fractionation associated with the enzy-
one radiogenic isotope (14C). Carbon is the main matic carboxylation step of carbon assimilation
constituent of the biosphere, rapidly recycled in in autotrophic organisms (Fig. 1). This fraction-
the ocean–atmosphere system. Carbon isotope ation is the main process responsible for the
ratios are easily measured, and therefore stable creation of isotopically light organic matter and C
carbon isotopes are widely used to trace the car- a remnant inorganic carbon pool that is isotopi-
bon cycle on Earth. Carbon isotopes may equally cally heavy relative to carbon sourced from
have utility in tracing the carbon cycle on other the Earth’s mantle, which has a d13C value of
planets. Stable carbon isotopes measured from ~5 ‰ (e.g., Broecker 1970). However, there is
samples collected in a geological context can be a wide range of abiotic and biotic processes
used to differentiate between biotic and abiotic (as discussed below) that cause significant car-
carbon transformations and can, therefore, serve bon isotope fractionations. Therefore, the pres-
as a biosignature. ence of an isotopically light or heavy carbon
pool is not a definitive evidence for autotrophic
carbon fixation.
Overview
Carbon isotopes are one of the most extensively Basic Methodology

used geochemical tracers of biological processes
on both local and global scales. Carbon isotopes There are several methods employed to measure
have been essential to the development of our carbon isotopes. The most standard technique
understanding of modern carbon fluxes and the involves CO2 analysis on a gas source isotope
long-term evolution of the global carbon cycle ratio mass spectrometer (IR-MS), using gas pre-
(e.g., Schidlowski 1988). pared with either combustion of organic carbon
Most of Earth’s carbon (~98.89 %) occurs as or acidification of inorganic carbon. Mass spec-
the lighter stable isotope (12C). The remaining trometers equipped with a dual inlet system
~1.1 % of Earth’s carbon occurs as the heavier require offline combustion or acidification and
(13C) stable isotope (de Laeter et al. 2003). There provide the highest precision, but standard devi-
are trace (<1 ppt) levels of radiogenic carbon ation on continuous flow IR-MS is commonly
(14C), which are generated in the upper atmo- <0.15 ‰. Accurate carbon isotope measure-
sphere through interaction between divalent ments can also be made using absorption spec-
nitrogen and background radiation. The half-life troscopy coupled with a diode laser (tunable
of 14C is 5,730 40 years (Scott 2003). Varia- diode laser absorption spectroscopy [TDLAS])
tions in carbon isotopes are reported in standard and cavity ring-down spectroscopy (CRDS). In
delta notation: situ techniques to analyze solid samples include
secondary ion mass spectrometry (SIMS) and
13 12 laser ablation inductively coupled plasma mass
ð C= CÞsa
d13 C ¼ 1 1000ð0=00, PDBÞ spectrometry (LA-ICP-MS). Both techniques
ð13 C=12 CÞst
offer high spatial resolution, on the order of a
few microns, and sub-per mil precision. An alter-
where the standard (st) is an international refer- native nondestructive technique is Raman spec-
ence material ([V]-PDB [Vienna Pee Dee troscopy, but analytical errors can be
Belemnite]). Carbon stable isotope ratios in nat- prohibitively large. Infrared spectroscopy is
ural samples range from about 100 ‰ to another emerging nondestructive method to mea-
+50 ‰. However, almost all natural samples sure carbon isotopes.
δ13C (‰, V-PDB)
−40 −30 −20 −10 0
Marine carbonate (past) > +10‰

Marine bicarbonate
Mantle carbon
Modern atmospheric CO 2
Modern dissolved CO2
Land plants
Eukaryotic algae
Cyanobacteria
Anoxygenic photosynthetic bacteria

Methanogenic
bacteria
−85‰ Methanotrophic bacteria
−54‰ Organic compounds formed by serpentinization reactions
−40 −30 −20 −10 0
Carbon Isotopes as a Geochemical Tracer, Fig. 1 Range of carbon isotope values in common inorganic and
biological carbon reservoirs (Modified from Schidlowski 2001)
Key Research Findings and Applications heterogeneity in the mantle; for example,
eclogites, which represent subducted oceanic
Carbon isotopes have been used extensively to crust, commonly possess more negative isotopic
trace changes in the global carbon cycle. The use values (Deines et al. 1991). Thus, the ratio of
of carbon isotopes to track the evolution of car- organic to carbonate carbon burial can be esti-
bon fluxes stems from the assumption that there mated using their isotopic compositions. Open-
must be an isotopic balance between the contin- marine carbonates typically have a d13C value of
ually renewed source of mantle carbon and the ~0 ‰, and average organic matter has a d13C
burial of organic and carbonate carbon value ~25 ‰. It follows that about 20 % of
(Schidlowski 2001). This relationship between carbon today is buried as organic matter
mantle-derived carbon (Cprim) and burial of (Holland 1978).
organic (Corg) and carbonate (Ccarb) carbon on This concept is interesting from an
the geological timescale is determined by the astrobiological perspective since carbon stable
relative burial rate of organic carbon (R) and isotope value can be used to track the evolution
can be expressed mathematically as of the carbon cycle through Earth’s history. The
isotopic composition of the Earth’s mantle is
generally assumed to have been essentially con-
d13 Cprim ¼ Rd13 Corg þ ð1 RÞd13 Ccarb
stant through time, in contrast to that of marine
carbonates and organic matter. However, the
The isotopic composition of the mantle flux of composition of both carbonates and organic mat-
inorganic carbon (d13Cprim) is, based on the carter can be measured at almost any point in Earth’s
bon isotope composition of diamonds and carbon history, since both these carbon pools are com-
dioxide released from volcanoes, generally mon in the geologic record. Importantly, the car-
accepted to be around ~5 ‰. However, carbon bon isotope composition of ancient carbonates is
isotope values of diamonds indicate some roughly equal to that of the dissolved inorganic
Carbon Isotopes as a Geochemical Tracer 369
Phanerozoic Proterozoic Archean

Neo- Meso- Paleo- Neo- Meso- Paleo-
δ13C (‰, V-PDB)
12
8
4
0 C
−4
0 500 1000 1500 2000 2500 3000 3500
Age (Ma)
Carbon Isotopes as a Geochemical Tracer, Fig. 2 Idealized record of the d13C values of marine carbonates through
time (Modified from Karhu 1999)
carbon reservoir, since only a small positive frac- remained essentially constant. This relative stasis
tionation takes place during mineral precipita- is a testament to the importance of stabilizing
tion. Therefore, the carbon isotope values of feedbacks at work in Earth’s biogeochemical
carbonates deposited in open-marine settings cycles. Short-lived (<5-million-year) isotope
usually reflect time-averaged seawater composi- excursions are relatively common in the Phanero-
tion. The d13C value of ancient organic matter is, zoic (the eon encompassing the last 542 million
similarly, usually roughly equivalent to that of years of Earth’s history). The magnitudes of these
organic matter when first sedimented (Hayes excursions are less than 5 ‰, and, in most cases,
et al. 1983). However, the composition of organic they can be linked to temporary increased or
matter is much more strongly affected by local decreased organic carbon burial. For instance,
factors such as rates of primary and secondary transient shifts to anoxic deep-sea conditions in
productivity, which are ultimately linked with the Cretaceous period resulted in burial of organic-
ocean redox state and chemical composition rich black shales and, consequently, positive car-
(e.g., Bekker et al. 2008) than is carbonate carbonate carbon isotope excursions (e.g., Sageman
bon. Therefore, carbon isotope studies of carbon- et al. 2006). Additionally, there are two major
ates and organic matter produced in open-marine periods in Earth’s history with long-lived carbon
settings hold the potential (when used with cau- isotope excursions. The mid-Paleoproterozoic
tion) to constrain past organic and carbonate (~2,300–2,100 million years ago) record is notable
burial fluxes, even if samples are billions of for an abundance of carbonates with markedly
years old. positive carbon isotope values, in cases reaching
Use of these methodologies has yielded a sur- even above 10 ‰ (Karhu and Holland 1996;
prising result: the d13C values of marine carbon- Bekker et al. 2003, 2008). In the Neoproterozoic
ates have remained relatively constant (~800–650 million years ago), markedly positive
throughout Earth’s history (Fig. 2). Almost all carbonate carbon isotope excursions are also com-
marine carbonates, dating back to 3.5 billion mon (Kaufman and Knoll 1996). These isotope
years ago, have a d13C value of ~0 ‰ signatures are traditionally interpreted as marking
(Schidlowski 1975). This indicates that even in periods of enhanced organic carbon burial. Car-
the Earth’s early history, there was significant bonates tracking the isotope composition of the
and continuous burial of isotopically light car- marine DIC reservoir that have markedly negative
bon, suggesting autotrophic primary production. d13C values are thought to reflect oxidation of a
Further, it implies that despite dramatic changes large organic carbon pool or extensive methane
to the biosphere, such as the evolution of land oxidation.
plants and calcareous phytoplankton, the relative Beyond tracking global processes, carbon iso-
burial of organic versus carbonate carbon has topes provide a means to trace local carbon
fixation pathways and carbon transformations in depleted simple organic molecules besides meth-
isolated systems. In contrast to the global carbon ane (e.g., Foustoukos and Seyfried 2004) and are
cycle, in isolated systems, the isotopic fraction- a potential source of abiogenic light organic car-
ations associated with abiogenic processes can be bon in the geologic record. Consequently, carbon
very important. For instance, the small isotopic isotopes are a powerful geochemical tracer, but
fractionation caused by carbon dioxide degassing there is often more than one explanation for the
during evaporation (~1 ‰) can create isotopi- d13C values. Therefore, whether the end goal is to
cally enriched dissolved inorganic carbon (DIC) trace the global carbon cycle, constrain carbon
pools, since evaporation in a closed system pathways in an isolated system, or find evidence
allows for expression of Rayleigh distillation of biological activity, a strong geological context
effects. Similarly, although the fractionation is an essential prerequisite to using carbon iso-
associated with carbonate mineral precipitation topes to derive a unique explanation.
is very small, in settings with efficient distillation
mechanisms, this process can form isotopically
distinct carbon pools. There are also ancient car- Future Directions
bonates, so-called diagenetic carbonates, that
record the isotopic composition of pore waters Carbon isotopes have greatly improved our
rather than the d13C value of DIC a large aquatic understanding of the carbon cycle on Earth. Car-
system. Pore water isotopic composition can be bon isotopes will undoubtedly play a significant
strongly influenced by organic matter reminera- role in unraveling aspects of the carbon cycle on
lization and thermal decarboxylation. other planets. With recent advances in in-flight
Methane cycling can produce markedly nega- technology, the idea of carbon isotope studies on
tive and positive carbon isotope values. Methane other planets is no longer a distant prospect in
on Earth is almost entirely directly or indirectly astrobiological studies.
biologically produced (Fiebig et al. 2009), fore-
most during methanogenesis, which can generate See Also
methane with d13C values <80 ‰. Although
the presence of isotopically light methane is com- ▶ Carbon Cycle, Biological
monly assumed to provide a signature of micro- ▶ Serpentinization
bial processes, abiogenic serpentinization can
also produce methane with highly negative d13C
values (Foustoukos and Seyfried 2004). Experi- References and Further Reading
mentally controlled serpentinization reactions
Beal EJ, House CH, Orphan VJ (2009) Manganese- and
can produce methane with d13C values <50 iron-dependent marine methane oxidation. Science
‰ (e.g., Horita and Berndt 1999). The presence 325:184–187
of d13Corg values as low as 60 ‰ in ca. 2,700- Bekker A, Karhu JA, Eriksson KA, Kaufman AJ
million-year-old sediments on several continents (2003) Chemostratigraphy of Paleoproterozoic carbonate
successions of the Wyoming Craton: tectonic forcing of
has been suggested to indicate aerobic methane biogeochemical change? Precambrian Res 120:279–325
oxidation (Hayes 1994; Eigenbrode and Freeman Bekker A, Holmden C, Beukes NJ, Kenig F, Eglington B,
2006) and may therefore date the rise of aerobic Patterson WP (2008) Fractionation between inorganic and
ecosystems. However, methane oxidation can organic carbon during the Lomagundi (2.22–2.1 Ga) car-
bon isotope excursion. Earth Planet Sci Lett 271:278–291
also be linked with ferric oxide, nitrate, or sulfate Bjerrum CJ, Canfield DE (2004) New insights into the
reduction (Orphan et al. 2001; Raghoebarsing burial history of organic carbon on the early Earth.
et al. 2006; Beal et al. 2009), and these oxidants Geochem Geophys Geosyst 5(8) (G3):2004GC000713
can be formed in an anoxic ocean–atmosphere Broecker WS (1970) A boundary condition on the evolution
of atmospheric oxygen. J Geophys Res 75:3553–3557
system. High temperature and pressure metal- de Laeter JR, Böhlke JK, De Bièvre P, Hidaka H, Peiser
catalyzed reactions (e.g., Fischer–Tropsch reac- HS, Rosman KJR, Taylor PDP (2003) Atomic weights
tions) can create a wide range of isotopically of the elements. Pure Appl Chem 75:683–800
Carbon Monosulfide 371
Deines P, Harris JW, Robinson DN, Gurney JJ, Shee SR MacKenzie FT, Lerman A (2006) Carbon in the
(1991) d13C and d18O variations in diamond and geobiosphere; Earth’s outer shell: topic in
graphite eclogites from Orapa, Botswana, and the Geobiology-25. Springer, New York, p 402
nitrogen content of their diamonds. Geochim Orphan VJ, House CH, Hinrichs K-U, McKeegan KD,
Cosmochim Acta 55:515–524 Delong EF (2001) Methane-consuming archaea
Eigenbrode JL, Freeman KH (2006) Late Archean rise of revealed by directly coupled isotopic and phylogenetic
aerobic microbial ecosystems. Proc Natl Acad Sci analysis. Science 293:484–487
103:15759–15764 Raghoebarsing AA, Pol A, van de Pas-Schoonen KT, Smol-
Fiebig J, Woodland AB, D’Alessandro W, Puttmann W ders AJ, Ettwig KF, Rijpstra WI, Schouten S, Damsté
C
(2009) Excess methane in continental hydrothermal JS, Op den Camp HJ, Jetten MS, Strous M (2006)
emissions is abiogenic. Geology 37:495–498 A microbial consortium couples anaerobic methane
Fischer WW, Schroeder S, Lacassie JP, oxidation to denitrification. Nature 440:878–879
Beukes NJ, Goldberg T, Strauss H, Horstmann UE, Rothman DH, Hayes JM, Summons RE (2003) Dynamics
Schrag DP, Knoll AH (2009) Isotopic constraints of the neoproterozoic carbon cycle. Proc Natl Acad Sci
on the Late Archean carbon cycle from the 100:8124–8129
Transvaal Supergroup along the western margin of Sageman BB, Meyers SR, Arthur MA (2006) Orbital time
the Kaapvaal Craton, South Africa. Precambrian Res scale and new C-isotope record for Cenomanian-
169:15–27 Turonian boundary stratotype. Geology 34:125–128
Foustoukos DI, Seyfried WE Jr (2004) Hydrocarbons in Schidlowski M (1988) A 3800-million-year isotopic
hydrothermal vent fluids: the role of chromium- record of life from carbon in sedimentary rocks.
bearing catalysts. Science 304:1002–1005 Nature 333:313–318
Hayes JM (1994) Global methanotrophy at the Schidlowski M (2001) Carbon isotopes as biogeochemical
Archean–Proterozoic transition. In: Bengtson S (ed) recorders of life over 3.8 Ga of Earth history: evolution
Early life on Earth (Nobel Symposium 84). Columbia of a concept. Precambrian Res 106:117–134
University Press, New York, pp 220–236 Schidlowski M, Eichmann R, Junge CE (1975) Precam-
Hayes JM (2001) Fractionation of the isotopes of carbon brian sedimentary carbonates: carbon and oxygen iso-
and hydrogen in biosynthetic processes. Rev Mineral tope geochemistry and implications for the terrestrial
Geochem 43:225–278 oxygen budget. Precambrian Res 2:1–69
Hayes JM, Waldbauer JR (2006) The carbon cycle and Scott EM (2003) The Fourth International Radiocarbon
associated redox processes through time. Philos Trans Intercomparison (FIRI). Radiocarbon 45:135–285
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Hayes JM, Kaplan IR, Wedeking KW (1983) Precambrian
organic geochemistry: preservation of the record. In:
Schopf JW (ed) Earth’s earliest biosphere: its origin
and evolution. Princeton University Press, Princeton, Carbon Monosulfide
pp 93–134
Holland HD (1978) The chemistry of the atmosphere and
oceans. Wiley, New York, p 351 William M. Irvine
Horita J, Berndt M (1999) Abiogenic methane formation Department of Astronomy, University of
and isotopic fractionation under hydrothermal condi- Massachusetts, Amherst, MA, USA
tions. Science 285:1055–1057
Hotinski RM, Kump LR, Arthur MA (2004) The effec-
tiveness of the Paleoproterozoic biological pump: a
d13C gradient from platform carbonates of the Pethei Synonyms
Group (Great Slave Lake Supergroup, NWT). Geol
Soc Am Bull 116:539–554 CS
Karhu J (1999) Carbon isotopes. In: Marshall CP,
Fairbridge RW (eds) Encyclopedia of geochemistry.
Kluwer, Dordrecht, pp 67–73
Karhu JA, Holland HD (1996) Carbon isotopes Definition
and the rise of atmospheric oxygen. Geology
24:867–870
Kaufman AJ, Knoll AH (1995) Neoproterozoic variations The diatomic molecule carbon monosulfide, CS, is
in the carbon isotopic composition of seawater: strat- widespread in interstellar and circumstellar
igraphic and biogeochemical implications. Precam- ▶ molecular clouds, in our ▶ Milky Way and in
brian Res 73:27–49 external galaxies. Its emission at millimeter wave-
Kump LR, Arthur MA (1999) Interpreting carbon-isotope
excursions: carbonates and organic matter. Chem Geol lengths is frequently used to estimate the density in
161:181–198 these regions, since these pure rotational
372 Carbon Monoxide
transitions require densities greater than about 105 Overview

molecules per cubic centimeter to be excited by
collisions. Various isotopic variants have been Carbon monoxide is not known to occur as a pure
observed astronomically, including those with substance in most planetary environments, but
13
C, 34S, and 33S, as well as the most abundant instead occurs primarily as a minor component
12 32
C S. CS is also observed in cometary comae. in gas and ice mixtures. Carbon monoxide is the
most abundant constituent after molecular hydro-
gen in interstellar space (see ▶ Molecules in
History Space), and spectroscopic observations indicate
it composes several percent of the icy component
Interstellar CS was first detected by radio astron- of ▶ comets (which is predominantly water ice).
omers in 1971. These observations indicate carbon monoxide
was a significant reservoir of carbon during for-
mation of the solar system. In planetary bodies,
See Also carbon monoxide primarily occurs as a trace
component in atmospheres (0.1 parts per mil-
▶ Comet lion on the present Earth) and in volcanic gases.
▶ Milky Way Transformations between carbon monoxide
▶ Molecular Cloud and ▶ carbon dioxide (CO2) proceed fairly read-
▶ Molecules in Space ily through photochemical reactions in planetary
▶ Sulfur atmospheres and through mechanisms such as the
so-called water-gas shift reaction in geologic
environments (CO + H2O $ CO2 + H2). The
latter is an oxidation-reduction reaction, and at
Penzias AA, Solomon PM, Wilson RW, Jefferts KB
the prevailing oxidation state of the Earth’s inte-
(1971) Interstellar carbon monosulfide. Astrophys rior, carbon dioxide is strongly favored by chem-
J 168:L53–L58 ical thermodynamics relative to carbon
monoxide. Consequently, CO2:CO ratios in vol-
canic gases on Earth are typically in a range of
Carbon Monoxide 103–105. Carbon monoxide may also be gener-
ated from dehydration of ▶ formic acid and vice
Thomas McCollom versa (HCOOH $ CO + H2O).
Laboratory for Atmospheric and Space Physics, Carbon monoxide readily forms complexes
University of Colorado, Boulder, CO, USA with transition metals, such as iron carbonyl
[Fe(CO)5] (carbonyl is the name given to the CO
radical). Formation of carbonyls from carbon
Synonyms monoxide can also occur on the surface of transi-
tion metal-bearing minerals and alloys. Carbonyls
CO are highly reactive, and carbonyls derived from
carbon monoxide have been hypothesized to be a
key reactant for the in situ formation of prebiotic
Definition organic compounds in several scenarios for the
▶ origin of life (e.g., Huber and Wächtershäuser
Carbon monoxide is a diatomic compound with 1997). One such reaction that is frequently
the chemical composition CO. At terrestrial invoked for the formation of organic matter in
atmospheric pressure, the melting point of pure the early solar system is Fischer-Tropsch synthe-
carbon monoxide is 205 C and the boiling sis, a surface-catalyzed process for conversion of
point is 191.5 C. gaseous mixtures of CO and H2 to hydrocarbons
Carbonaceous Chondrite 373
and other functionalized organic compounds such sustaining life as we know it and the main ele-
as fatty acids. Potential environments for the ment in all classes of macromolecules. All cells
surface-catalyzed conversion of carbon monoxide require carbon as a major nutrient. On a dry
to prebiotic organic compounds by the Fischer- weight basis, a typical cell is 50 % carbon.
Tropsch synthesis or other processes include dust Depending on their carbon source, organisms
grains in the early solar system, volcanic fuma- are either heterotrophs, requiring one or more
roles, and hydrothermal systems. organic compounds as their carbon source, or C
autotrophs, where CO2 is the carbon source.
See Also
See Also
▶ Carbon Dioxide
▶ Comet ▶ Biosynthesis
▶ Fischer-Tropsch-Type Reaction ▶ Carbon Cycle, Biological
▶ Formic Acid ▶ Carbon Dioxide
▶ Interstellar Ices ▶ Chemotroph
▶ Interstellar Medium ▶ Macronutrient
▶ Origin of Life ▶ Metabolism
▶ Prebiotic Chemistry ▶ Phototroph
Huber C, Wächtershäuser G (1997) Activated acetic acid Carbonaceous Chondrite

by carbon fixation on (Fe, Ni)S under primordial con-
ditions. Science 276:245–247 Tilman Spohn and Frank Sohl
Deutsches Zentrum f€ur Luft- und Raumfahrt
(DLR), Institut f€ur Planetenforschung, Berlin,
Germany
Carbon Nitride
▶ Cyanogen Synonyms
Carbonaceous meteorite; CC
Carbon Source
Definition
Felipe Gomez
Centro de Astrobiologı́a (CSIC/INTA), Instituto Carbonaceous chondrites constitute a
Nacional de Técnica Aeroespacial, Torrejón de subcategory of ▶ chondrites – which in turn are
Ardoz, Madrid, Spain stony ▶ meteorites. Carbonaceous chondrites are
the most primitive meteorites identified thus far
and are mostly regarded as remnants of the first
Definition solid bodies to accrete in the ▶ solar nebula. The
main components of carbonaceous chondrites are
Carbon source refers to any carbon-containing ▶ chondrules and ▶ CAIs (Ca-Al-rich inclu-
molecule (carbohydrate, amino acid, fatty acid, sions), which are embedded in a matrix of
CO2) used by an organism for the synthesis of its micrometer-sized dust particles. Since carbona-
organic molecules. Carbon is a basic element for ceous chondrites contain the highest
374 Carbonaceous Chondrites, Organic Chemistry of
concentration of volatile elements and molecules enough to undergo planetary differentiation and
(e.g., ▶ organic refractory matter) of the chon- accreted as aggregate rocks. CC have varying but
drites, they are thought to have formed at the near solar chemical composition and contain carbon
lowest temperatures. Their chemical composition in various percent amounts as well as water and all
is very similar to that of the Sun (albeit depleted other biogenic elements, HONPS. Most of CC car-
in ▶ hydrogen and helium), and thus they can be bon is in organic materials found both as kerogen-
considered (apart from ▶ comets) to be the most like insoluble macromolecules and soluble organic
primitive material present in our ▶ solar system. compounds, such as amino acids; the relative pro-
portions of these organic components change with
classification and the bulk water composition may
See Also also vary from over 20 % (CI) to less than 1 % (CV).
Meteorites are generally categorized as falls or finds
▶ CAIs depending on whether their falls had been observed
▶ Chondrite or not. CC have been classified, so far, in eight
▶ Chondrule group types: CI, CM, CR, CK, CH, CB, CO, CV,
▶ Comet where the second letter refers to the first meteorite
▶ Meteorites classified for the type or a compositional character-
▶ Solar Nebula istic (respectively: Ivuna, Mighei, Renazzo,
▶ System Solar Formation, Chronology of Karoonda, High metal, Bencubbin, Ornans,
Vigarano). Some rare CC of recent falls, such as
Tagish Lake and Sutters’ Mill remain unclassified
or have been simply classified as C. Falls are more
Carbonaceous Chondrites, Organic desirable for organic analyses since they have been
Chemistry of less exposed to terrestrial contamination. Both Jap-
anese and American institutions have established
Conel Michael O’Donel Alexander1 and programs for the search of meteorites in Antarctica
Sandra Pizzarello2 during which many pristine samples have been
1
Department of Terrestrial Magnetism, Carnegie collected thanks to their preservation in the glaciers
Institution of Washington, NW Washington, DC, (http://mineralsciences.si.edu/research/meteorites/
USA antarctica/).
2
Department of Chemistry & Biochemistry,
Arizona State University, Tempe, AZ, USA
Overview
Keywords CCs are primitive stony meteorites of elemental

composition very similar to that of the Sun that
Amino acids; Asteroids; Carbonaceous; have the distinction of containing carbon mostly
Chondrites; Comets; Enantiomeric excess; Inter- as organic materials and offer a so far unique
planetary dust particles; Interstellar medium; record of organic chemistry in the Solar System
Isotope anomalies; Meteorites; Molecular cloud; at the time the Earth was formed and before the
Nucleic acids; Organic matter; Protoplanetary inception of life. CC are heterogeneous in com-
disk position, with their carbon varying between types
in both total content, ~1.5–4 %, and relative dis-
tribution of organic materials, which are classi-
Definition fied procedurally as soluble compounds or
insoluble organic material (IOM).
Carbonaceous Chondrites (CC) are stony fragments The compounds found so far in CC comprise:
of primitive asteroids that never became large water-soluble carboxylic-, dicarboxylic-, amino-,
Carbonaceous Chondrites, Organic Chemistry of 375
hydroxy-, keto-, and pyridine carboxylic-acids; Although the origins of CC soluble and insol-
sugar alcohols and acids, aldehydes and ketones; uble materials have yet to be established, both are
amines and amides; purine bases and pyrimidine likely to be the product of a long cosmic history,
basesand solvents-soluble aliphatic, aromatic and which is revealed by their isotopic composition
hetero-atom containing hydrocarbons. Several showing enrichments of 2H (Deuterium), 15N and
13
CC compounds have counterparts in the bio- C abundances that point to synthesis in very
sphere, e.g., 12 amino acids are found in terres- cold environments, e.g., some CR amino acids C
trial proteins: glycine, alanine, proline, valine, have dD values of over 300 ‰ as measured in the
leucine, isoleucine, serine, threonine, tyrosine, Interstellar medium (ISM). The current under-
phenylalanine, aspartic- and glutamic acids standing for the formation of CC organic com-
(Pizzarello and Holmes 2009). pounds is based on data for their isotopic
Another similarity between terrestrial and fractionation, detections of organic compounds
meteoritic amino acids is their chirality and the in the ISM, the extensive presence of water in
occurrence in some of L-enantiomeric excesses asteroidal parent bodies recorded by the mineral
(Pizzarello and Groy 2011). This chiral asymme- phases and is known as “the interstellar-parent
try involves so far only few CC amino acids, is body formation hypothesis”. It has been proposed
not constant for the same amino acid in different that interstellar precursors molecules were incor-
types of meteorites and is of unknown origin. porated in asteroid and further processed by aque-
Nevertheless, it shows the presence in abiotic ous reactions into the various molecular species
environments of possible chiral influences and extracted from meteorites.
enhances the astrobiological appeal of meteoritic
compounds. Contrary to the compositional selec-
tivity and homochirality of biomolecules, how- See Also
ever, most meteoritic compounds do not show
chiral preferences and are found with almost ▶ Aliphatic Hydrocarbon
complete structural diversity for a given carbon ▶ Amino Acid
number, e.g., close to 100 amino acids are recog- ▶ Aromatic Hydrocarbon
nized in meteorite extracts compared to the ▶ Asteroid
23 that are constituents of terrestrial proteins ▶ Comet (Nucleus)
(Pizzarello and Shock 2010). ▶ Enantiomeric Excess
The IOM is a structurally complex, macromo- ▶ Globule, Nanoglobule
lecular material with a bulk elemental composition ▶ Interplanetary Dust Particle
of C100H79N4O15S3, which is similar to the aver- ▶ Interstellar Dust
age of comet Halley CHON particles, and is dis- ▶ Interstellar Medium
persed in chondritematrices as particles that are ▶ Meteorite, Murchison
typically <1 mm across; it is also compared often ▶ Meteorites
to terrestrial kerogens for its content of S and ▶ Protoplanetary Disk
P besides H, C, O, N, insoluble nature and hetero- ▶ Racemization
geneity. The IOM is likely the result of several ▶ Serpentinization
chemical processes and, like kerogens, a mix
aggregate of various phases and functional groups
as indicated by pyrolytic and spectroscopic studies References and Further Reading
(Alexander et al. 2007). The complexity of the
IOM is further displayed by the materials’ suscep- Alexander CMO’D, Fogel M, Yabuta H, Cody GD
tibility to release water- and solvents-soluble com- (2007) The origin and evolution of chondrites recorded
in the elemental and isotopic compositions of their
pounds, such as ammonia and hydrocarbons under
macromolecular organic matter. Geochim Cosmochim
hydrothermal conditions (300 C, 100MP) Acta 71:4380–4403
(Pizzarello et al. 2011; Yabuta et al. 2007). http://mineralsciences.si.edu/research/meteorites/antarctica/
376 Carbonaceous Meteorite
Pizzarello S, Groy TL (2011) Molecular asymmetry in (ZnCO3), and cerussite (PbCO3), which result from
extraterrestrial organic chemistry: an analytical per- surficial alteration of metallic ore deposits. The
spective. Geochim Cosmochim Acta 75:645–656.
doi:10.1016/j.gca.2010.10.025 term is also applied to sediments or ▶ sedimentary
Pizzarello S, Holmes W (2009) Nitrogen-containing com- rocks such as limestones or dolostones that are
pounds in two CR2 meteorites: 15N composition, composed predominantly of carbonate minerals.
molecular distribution and precursor molecules. In the Precambrian, particularly the Proterozoic,
Geochim Cosmochim Acta 73:2150–2162.
doi:10.1016/j.gca.2009.01.022 carbonate minerals precipitated on microbial mats
Pizzarello S, Shock E (2010) Perspectives in biology. by their photosynthetic activity and resulting alka-
Cold Spring Harbor Laboratory Press, pp 89–107. linity of the surface are common. Most Phanero-
doi:10.1101/cshperspect.a002105 zoic carbonates are composed to a large degree of
Pizzarello S, Williams BL, Lehman J, Holland PH, Yarger
JL (2011) Abundant ammonia in primitive asteroids the shells, tests, and spicules or their fragments of
and the case for a possible exobiology. Proc Natl Acad marine organisms, cemented by secondary carbon-
Sci USA 108:4303–4306. www.pnas.org/cgi/doi/ ates. Rarely, carbonates precipitate chemically
10.1073/pnas.1014961108 directly from sea- or lake water. Dolomite, a wide-
Yabuta H, Williams LB, Cody GD, Alexander CMO’D,
Pizzarello S (2007) The insoluble carbonaceous mate- spread carbonate, has variable origins, but its pre-
rial of CM chondrites: a possible source of discrete cipitation is also known to be biologically
organic compounds under hydrothermal conditions. mediated. Carbonate sediments are rare in Archean
Meteorit Planet Sci 42:37–48. http://meteoritics.org sequences but common as cements.
Carbonate minerals detected by Martian
rovers may provide evidence of the presence of
Carbonaceous Meteorite water of the surface early in the history of
▶ Mars.
▶ Carbonaceous Chondrite
▶ Meteorite, Murchison
See Also
▶ Mars
Carbonate ▶ Sedimentary Rock
Nicholas Arndt
Carbonate, Extraterrestrial
Synonyms Daniela Tirsch

German Aerospace Center DLR, Institute of
Calcareous sediment; Limestone Planetary Research, Berlin, Germany
The term carbonate either refers to a mineral or to a The term carbonate can either refer to carbonate
rock. Examples of carbonate minerals are calcite minerals, which contain the carbonate ion CO2 3 ,
(CaCO3) and dolomite (MgCa(CO3)2), which are or to carbonate rocks, which contain carbonate
common constituents of limestones and other cal- minerals. On Earth, the vast majority of carbon-
careous sediments; siderite (FeCO3), which also ate rocks are of sedimentary origin, while igneous
occurs in ▶ sedimentary rocks; magnesite carbonates (carbonatites) are extremely rare. Fur-
(MgCO3), an alteration product of ultramafic thermore, the formation of carbonate minerals on
rock; and malachite Cu2(CO3)(OH)2, smithsonite Earth generally takes place within an aqueous
Carbonyl 377
solution, either through abiotic precipitation or reaction (HCO3 ) keeps the pH of the ocean high
biomineralization. Hence, the presence of car- enough for the carbonate ion to be abundant and
bonates on other planets (especially Mars) is enhances the precipitation of calcium carbonates.
viewed as an indicator for the former presence
of liquid water. Extraterrestrial carbonates have
See Also
been identified in a wide range of localities on
Mars as well as in several Martian meteorites. C
▶ Weathering
They include magnesium carbonate detections
in the Nili Fossae region, carbonates intermixed
with iron/magnesium-smectites (phyllosilicates)
in the Libya Montes region, magnesium-rich car-
bonates at Gusev crater, and calcium-/iron-rich Carbonization
carbonates at various spots in Tyrrhena Terra.
The spectral signatures share similarities with ▶ Pyrolysis
carbonatious chondrites.
See Also Carbonyl
▶ Mars Henderson James (Jim) Cleaves II

▶ Mineral Earth–Life Science Institute (ELSI), Tokyo
▶ Phyllosilicates, Extraterrestrial Institute of Technology, Meguro–ku, Tokyo,
▶ Sulfates, Extraterrestrial Japan
Washington, DC, USA
Carbonation
Francis Albarède
Ecole Normale Supérieure de Lyon, Lyon,
France
Definition
In organic chemistry, a carbonyl is a functional

group consisting of a carbon atom doubly bonded
Definition
to an oxygen atom (Fig. 1). Some examples of
carbonyl-group-containing compounds include
A carbonation reaction describes the breakdown
carboxylic acids and their derivatives (e.g.,
of silicates by dissolved carbon dioxide. For
amides, esters, and anhydrides), aldehydes, and
example, the reaction of olivine carbonation
ketones. The term carbonyl can also refer to a
reads
ligand in an organometallic complex (e.g., iron
pentacarbonyl, Fe(CO)5).
Mg2 SiO4 ðsÞ þ 4CO2 ð1Þ þ 2H2 O
Since oxygen is more electronegative than
¼ 2Mg2þ ð1Þ þ 4HCO3 ð1Þ þ SiO2 ðsÞ carbon, the oxygen atom in a carbonyl group
pulls electron density toward itself and away
Carbonation enhances the drawdown of volcanic from carbon making the bond polar. The carbonyl
CO2 by ▶ weathering; it thus limits the partial carbon thus tends to be electrophilic and more
pressure PCO2 and therefore the greenhouse reactive with nucleophiles. The electronegative
effect. In addition, the alkalinity liberated in the oxygen can react with an electrophile.
378 Carbonyldiamide
Carbonyl, Fig. 1 General O carboxylic acids of biological importance, for

structure of carbonyl example, fatty acid esters are important compo-
compounds
nents of many cell membranes, proteins are poly-
R1 R2 mers of amino acids, and many compounds in
intermediary metabolism are carboxylic acids.
Common prebiotic syntheses of carboxylic acids
Protons bonded to carbon atoms alpha to a
proceed via the hydrolysis of precursor nitriles.
carbonyl group are often considerably more
Because they are both hydrogen-bond accep-
acidic (by 3 pKa units) than typical aliphatic
tors and hydrogen-bond donors, they are able to
C-H bonded protons. The carbonyl group is in
participate in hydrogen bonding. Carboxylic acids
tautomeric equilibrium with an enol configura-
tend to have higher boiling points than water partly
tion. Deprotonation of this enol produces an
because of their tendency to form hydrogen-
enolate anion, which is nucleophilic, and can
bonded dimers. Carboxylic acids are polar: short-
alkylate electrophiles such as other carbonyls.
chain aliphatic carboxylic acids (e.g., those
containing one to five carbon atoms) are soluble
in water, whereas higher carboxylic acids are less
See Also
soluble due to the increasingly hydrophobic nature
of their aliphatic components.
▶ Aldehyde
Carboxylic acids form various derivatives in
combination with other functional group-
containing compounds, for example, esters in
combination with alcohols, amides in combina-
Carbonyldiamide tion with amines, and anhydrides in combination
with another carboxylic acid.
▶ Urea
See Also
Carboxylic Acid ▶ Acetic Acid

▶ Amide
Henderson James (Jim) Cleaves II ▶ Ester
Earth–Life Science Institute (ELSI), Tokyo ▶ Formic Acid
Institute of Technology, Meguro–ku, Tokyo, ▶ Nitrile
Japan
Washington, DC, USA Carboxylic Acids, Geological
Center for Chemical Evolution, Georgia Institute Record of
Jennifer Eigenbrode
Definition NASA Goddard Space Flight Center, Greenbelt,
MD, USA
In chemistry, carboxylic acids are organic acids
that contain a carboxyl functional group. The gen-
eral formula of a carboxylic acid is R-COOH. Keywords
Carboxylic acids are proton donors. Some com-
mon examples are ▶ formic acid, H-COOH, and Carbonyl; Fatty acids; Hydroxyl; Organic acid;
▶ acetic acid, CH3COOH. There are many Carboxyl; Lipids
Carboxylic Acids, Geological Record of 379
Synonyms regions in the same molecule. The amphiphilic

nature of these molecules leads to the formation
Fatty acids of nonbiological monolayers on the water and
mineral surfaces or micelles, sphere-shaped clus-
ters, in solution. Biology takes advantage of the
Definition amphiphilic nature of carboxylic acids to form
fatty acid lipid bilayers of cellular ▶ membranes. C
▶ Carboxylic acids are a class of organic com- Formation of nonbiological monolayers was
pounds that contain one or more carboxyl groups likely critical for the formation of the earliest
per molecule. Each carboxyl group, which is a cellular life.
combination of ▶ carbonyl and hydroxyl groups, Carboxylic acids are weak acids that only par-
has the formula Cð¼ OÞOH ðor COOHÞ . tially dissociate into H+ cations and RCOO-
Carboxylic acids are polar and proton donors. anions in pH neutral aqueous solution. In the
Carboxylic acids are widespread in nature, often presence of a base, carboxylic acids become car-
combined with other functional groups, and ubiq- boxylates. These moieties are important in geo-
uitous in biology, recent sediments, and carbona- logical and biogeochemical processes because
ceous meteorites. As they degrade during they can be siderophores, chelating with metals,
diagenesis and thermal maturation in the rock such as ferric iron.
record, they generally transform into esters, car- The carbon atom of a carboxyl group is in a
boxylates (i.e., hydroxyl group replaced with relatively high ▶ oxidation state and can be par-
salts or anions), and, eventually, alkanes. tially reduced by sulfur or other organic mole-
cules during diagenesis under reducing
conditions. ▶ Oxidation of the carboxyl carbon
commonly occurs in oxidizing environments to
Overview form carbon dioxide (decarboxylation reaction).
Depending on the geochemical or biological
Carboxyl groups are polar. The carbonyl group is reaction, the alkyl side chain may end up reduced
a hydrogen-bond acceptor, and the hydroxyl to an alkane or oxidized, often with the addition
group is a hydrogen-bond donor. Consequently, of an anion or salt. In both cases, decarboxylation
carboxylic acids participate in hydrogen bonding, results in the loss of a carbon from the carbon
particularly with each other and with minerals skeleton, and the product can then be preserved in
such as phyllosilicates, oxyhydroxides, and the rock record.
other oxides. In addition, the “self-association”
of carboxylic acids that results from hydrogen
bonding generates stabilized dimeric pairs in
nonpolar media. The dimers have decreased vol- See Also
atility. The bonds must be broken chemically or
thermally for the carboxyl group to be reactive to ▶ Acid Hydrolysis
other geochemicals. The stabilization of dimers ▶ Carbonaceous Chondrites, Organic
and the nonreactive nature of the alkyl side chain Chemistry of
(or other carbon skeleton) contribute to the ▶ Carbonyl
refractory nature of carboxylic acids during dia- ▶ Carboxylic Acid
genesis compared to more polar organic ▶ Cell Wall
molecules. ▶ Complex Organic Molecules
Carboxylic acids having more than five car- ▶ Fatty Acids, Geological Record of
bons in an alkyl chain are not soluble in water ▶ Kerogen
unless at high temperatures and exhibit both ▶ Membrane
hydrophilic (carboxyl) and hydrophobic (alkyl) ▶ Oxidation
380 Carboxysomes, Structure and Function
References and Further Reading is to concentrate CO2 with close proximity to

RuBisCO, eliminating the competitive reaction
Eigenbrode JL (2007) Fossil lipids for life-detection: a with oxygen. Thus, carboxysomes are the site of
case study from the early earth record. Space Sci Rev
carbon dioxide (CO2) fixation.
135:161–185
Kraemer SM, Butler A, Borer P, Cervini-Silva J (2005)
Siderophores and the dissolution of iron bearing min-
erals in marine systems. Rev Mineral Geochem History
59:53–76
Kraemer SM, Crowley D, Kretzschmar R (2006)
Siderophores in plant iron acquisition: geochemical In 1956 the first carboxysomes (then termed poly-
aspects. Adv Agron 91:1–46 hedral bodies) were visualized in a cyanobacterium
Sephton MA (2005) Organic matter in carbonaceous (Drews and Niklowitz 1956). Following the discov-
meteorites: past, present and future research. Phil
ery of similar structures in several other
Trans R Soc A 363:2729–2742
Wade LG (2009) Organic chemistry, 7th edn. Prentice- cyanobacteria and chemoautotrophs, carboxysomes
Hall, New Jersey, p 1320 were purified in 1973 and shown to contain the
enzyme, ribulose-1,5-bisphosphate carboxylase/
oxygenase (RuBisCO) (Shively et al. 1973).
Because RuBisCO plays a central role in carbon
Carboxysomes, Structure and fixation, they were given the name carboxysomes.
Function
Jeffrey Blanchard1 and Farah Abdul-Rahman2 Overview

1
Biology Department, University of
Massachusetts Amherst, Amherst, MA, USA Carboxysomes are intracellular organelles com-
2
University of Massachusetts Amherst, Amherst, posed of a proteinaceous shell that encapsulates
MA, USA the enzyme carbonic anhydrase and ribulose-1,5-
bisphosphate carboxylase/oxygenase (RuBisCO)
(Cameron et al. 2013). Unlike other types of
Keywords organelles, no lipids are associated with
carboxysomes. Carboxysomes are icosahedral in
Carbon dioxide; Calvin Cycle; Ribulose-1,5- shape, akin to bacterial phages and capsid
bisphosphate carboxylase/oxygenase viruses. However, there is no other evidence of
an evolutionary connection between
carboxysomes and any known viruses.
Synonyms X-ray crystallography of individual shell pro-
teins revealed hexamers as the basic microcom-
Bacterial microcompartments; Carbon concen- partment shell structure building block and
trating mechanisms; Polyhedral bodies showed how these hexamers assemble to form
the walls of polyhedral shell (Kerfeld
et al. 2005). Subsequent structural studies deter-
Definition mined that another universally conserved shell
protein forms pentameric structures that are
Carboxysomes are intracellular proteinaceous located in the vertices of the icosahedra (Tanaka
compartments found in cyanobacteria and et al. 2008). These structural studies have led to
some ▶ chemoautotrophs that encapsulate the beautiful structural models of carboxysomes that
enzymes carbonic anhydrase and ribulose-1,5- suggest mechanisms for the passage of molecules
bisphosphate carboxylase/oxygenase through the pores formed by hexameric and
(▶ RuBisCO). The function of the carboxysome pentameric structures (Yeates et al. 2008).
Carboxysomes, Structure and Function 381
Carboxysomes fall under the broader class of normal atmospheric CO2 levels and can only
structures, the bacterial microcompartments, grow in high (5 %) CO2 (Ogawa et al. 1994).
which encompasses several functional types. The ability of cyanobacteria to grow without
Bacterial microcompartments share evolution- carboxysome shells at higher CO2 concentrations
arily related shell proteins which encapsulate dif- suggests that these structures would not be
ferent enzymes. Other types of bacterial required if the atmosphere’s current makeup had
microcompartments have been shown to utilize higher CO2 levels. C
1,2-propanediol, ethanolamine, and choline
(Abdul-Rahman et al. 2013).
Carboxysomes are the site of carbon dioxide See Also
(CO2) fixation. Bicarbonate (HCO3) is
transported into the shell through nanopores, ▶ Oxygenation of the Earth’s Atmosphere
where the shell-associated enzyme, carbonic
anhydrase, converts HCO3 to CO2. RuBisCO
packed inside the carboxysome converts the sub- References and Further Reading
strates CO2 and ribulose-1,5-bisphosphate into
Abdul-Rahman F, Petit E, Blanchard JL (2013) The dis-
two 3-phospho-D-glycerate (3-PG) molecules. 3-
tribution of polyhedral bacterial microcompartments
PG then exits the carboxysome and can be used as suggests frequent horizontal transfer and operon
a metabolic intermediate in the Calvin-Benson- reassembly. J Phylogenet Evol Biol 1:e118
Bassham cycle. Cameron JC, Wilson SC, Bernstein SL, Kerfeld CA
(2013) Biogenesis of a bacterial organelle: the
RuBisCO has low affinity for CO2 and is
carboxysome assembly pathway. Cell
capable of binding O2. The oxygenation of 155(5):1131–1140
ribulose-1,5-bisphosphate produces Drews G, Niklowitz W (1956) Cytology of cyanophycea.
phosphoglycolate which can accumulate to toxic II. Centroplasm and granular inclusions of
Phormidium uncinatum. Arch Mikrobiol (in German)
levels and inhibit photosynthesis (Yeates
24(2):147–162
et al. 2008). Thus, the shell serves to both con- Kerfeld CA, Sawaya MR, Tanaka S, Nguyen CV,
centrate CO2 around RuBisCO and block oxygen Phillips M, Beeby M, Yeates TO (2005) Protein struc-
from enzymatically competing with CO2. tures forming the shell of primitive bacterial organ-
elles. Science 309(5736):936–938
Two types of carboxysomes have been char-
Ogawa T, Marco E, Orus MI (1994) A gene (ccmA)
acterized, the a- and b- carboxysomes, named for required for carboxysome formation in the cyanobac-
the form of RuBisCO they encapsulate. The terium Synechocystis sp. Strain PCC6803. J Bacteriol
b-carboxysome is found exclusively in 176(8):2374–2378
Session AL, Doughty DM, Welander PV, Summons RE,
cyanobacteria, while the a-carboxysome is
Newman DK (2009) The continuing puzzle of the
found in marine cyanobacteria as well as in great oxidative event. Curr Biol 19(14):R567–R574
some autotrophic ▶ proteobacteria. The Shively J, Ball F, Brown D, Saunders R (1973) Functional
carboxysome’s presence in all cyanobacteria, organelles in prokaryotes: polyhedral inclusions
(carboxysomes) of Thiobacillus neapolitanus. Science
with the exception of Cyanobacterium UCYN-A
(New York, NY) 182:584–586
which lacks genes for carboxysomes and the Cal- Tanaka S, Kerfeld CA, Sawaya MR, Cai F, Heinhorst S,
vin cycle (Tripp et al. 2009), indicates it is an Cannon GC, Yeates TO (2008) Atomic-level models
essential structure for CO2 fixation in the phylum. of the bacterial carboxysome shell. Science
319(5866):1083–1086
The great oxidation event (~2.4 billion years
Tripp HJ, Bench SR, Turk KA, Foster RA, Desany BA,
ago) altered the atmosphere with the appearance Niazi F, Affourtit JP, Zehr JP (2009) Metabolic
of free atmospheric oxygen and is attributed to streamlining in an open-ocean nitrogen-fixing cyano-
the emergence of photosynthetic cyanobacteria bacterium. Nature 464(7285):90–94
Yeates TO, Kerfeld CA, Heinhorst S, Cannon GC, Shively
that release oxygen as a by-product (Sessions JM (2008) Protein-based organelles in bacteria:
et al. 2009). Cyanobacterial mutants that lack carboxysomes and related microcompartments. Nat
one or more shell proteins cannot survive under Rev Microbiol 6(9):681–691
382 Cassini
Giovanni Domenico ▶ Cassini in 1675 and can

Cassini be observed with a small telescope. It extends
between 117,500 and 122,000 km from the center
Therese Encrenaz of Saturn. Observations by the Cassini orbiter
LESIA, Observatoire de Paris - Section de have shown that the Cassini division is not devoid
Meudon, Meudon, France of matter, as previously thought, but it consists of
a region where the density of particles is lower
than that in the main rings themselves. The exis-
Definition tence of the Cassini division is probably due to a
gravitational interaction with ▶ Mimas, as the
Giovanni Domenico Cassini (or Jean-Dominique Cassini division is in resonance with the orbit of
Cassini) (1625–1712) was born in Perinaldo, near this satellite. There are other divisions in Saturn
Naples, and was Professor at Bologna. In 1669, he rings (Encke division, Keeler division).
became a member of the French Académie des
Sciences, and in 1671, he became the first Director
of the Observatoire de Paris, from which the Paris See Also
Meridian was defined. Its first scientific aims were
metrology, celestial mechanics, and positional ▶ Mimas
astrometry. Cassini made important discoveries ▶ Planetary Rings
regarding solar-system objects: determination of ▶ Saturn
Mars’ rotation, Jupiter’s rotation, Jupiter’s satel-
lites’ revolutions, identification of several of Sat-
urn’s satellites (Iapetus, Rhea, Thetys, and Dione),
as well as the Cassini Division within Saturn’s Cassini Mission
rings. Cassini also recorded observations of the
▶ zodiacal light. The Cassini space mission, ▶ Cassini-Huygens Space Mission
launched in 1987 and presently exploring the
▶ Saturn system, was named after him.
Cassini Spacecraft
See Also
▶ Cassini-Huygens Space Mission
▶ Planetary Rings
▶ Saturn
Cassini State
Cassini Division Rory Barnes

Astronomy Department, University of
Therese Encrenaz Washington, Seattle, WA, USA
Meudon, Meudon, France
Definition
Definition When an orbiting body is affected by tides, its

spin axis, spin rate, and orbital plane can reach an
The Cassini Division is a gap between the A and equilibrium state in which obliquities (tilts)
B rings of ▶ Saturn. It was first identified by are nonzero. Cassini states are observed in the
Cassini-Huygens Space Mission 383
Earth-Moon system, Triton-Neptune system, and

possibly, Europa-Jupiter system. In these cases, Cassini-Huygens Space Mission
the satellite’s obliquity is locked at a small non-
zero value, but in general, with large inclinations, Athena Coustenis
the obliquity could be large. Laboratoire d’Etudes Spatiales et
d’Instrumentation en Astrophysique (LESIA),
Observatoire de Paris, CNRS, UPMC Univ. Paris C
See Also 06, Univ. Paris-Diderot, Meudon Cedex, France
▶ Tides, Planetary
Keywords
Cassini-Huygens; Cassini orbiter; Huygens

probe; Kronian satellites; Rings; Saturn; Titan;
Cassini Titan’s Probe Enceladus
▶ Huygens Probe
Synonyms
Cassini mission; Cassini spacecraft; Huygens

Cassini, Giovani Domenico probe
Fernando B. Figueiredo
CITEUC, University of Coimbra, Coimbra, Definition
Portugal
Cassini-Huygens is a large ▶ NASA-ESA-▶ ASI
mission, composed of the ▶ Saturn orbiter
History (▶ Cassini) and the ▶ Titan lander (▶ Huygens).
Cassini-Huygens reached Saturn in 2004 after a
Cassini, Giovanni Domenico [Jean-Dominique] trip that lasted about 7.5 years and has since then
(1625–1712) was an Italian astronomer born in been investigating the Saturnian environment,
Perinaldo in June 8, 1625. In 1669, Cassini set carrying out the first detailed survey of the
out for Paris (in 1673, became a naturalized ▶ planet, its rings, and the 62 currently known
Frenchman) where he will be appointed the first satellites, with a focus on ▶ Titan. The Cassini
director of the Observatory of Paris, starting that instruments have returned a great amount of data
way a family dynasty of prominent astronomers in that have revolutionized our view of the Satur-
prerevolutionary France. His astronomical works nian system. The orbiter is foreseen to remain
ranged from studies of comets (that he considered operational until 2017.
analogous to planets but traveling in paths of
greater eccentricity) to the movements of Jupiter’s
moons and Saturn’s satellites and ring. Cassini History
found the Saturnian satellites, Iapetus (1671),
Rhea (1672), and Tethys and Dione (1684) and An ambitious and international mission to
noted the division of the rings of Saturn. explore the Saturnian system was initially pro-
In 1997 in a joint effort between NASA, ESA, posed to the National Aeronautics and Space
and ASI, the Cassini Mission was launched to Administration (NASA) and the European
study the structure and physical properties and Space Agency (ESA) in 1982 by a team of Euro-
dynamics of Saturn and its satellites, mainly Titan. pean and US scientists. After extensive
384 Cassini-Huygens Space Mission
discussions between ESA and NASA, the initial One particular target of Cassini-Huygens was
concepts eventually evolved by 1989 into Titan, also visited in situ by the ▶ Huygens
Cassini-Huygens, a mission composed of an probe: on December 25, 2004, the Huygens
American orbiter and a European descent probe. probe separated from the orbiter and reached
This made it the first truly international planetary Titan on January 14, 2005, when it descended
mission, in addition to its other unique features. through the atmosphere, landed on the moon’s
The Italian Space Agency (ASI) is responsible for surface, and relayed scientific information for a
the spacecraft’s radio antenna and portions of total of more than 5 h (Lebreton et al. 2008).
three scientific instruments. Cassini-Huygens Mission control activities for Cassini are
arrived in the Saturnian system in July 2004. conducted from the Space Flight Operations
The Huygens probe executed its mission in Jan- Facility at the Jet Propulsion Laboratory
uary 2005. After two extensions, the Cassini (▶ JPL), where the project is headquartered.
orbiter continues its exploration of the Saturnian
system until 2017. For a thorough overview of the The Spacecraft and Its Payload
mission and its objectives, see “The Cassini- The spacecraft, including the orbiter and the
Huygens Mission” (Russell 2004). probe, is the largest and most complex
interplanetary spacecraft built to date. The orbiter
has a mass of 2,150 kg (plus fuel), and the probe
Overview has a mass of 350 kg. The Cassini spacecraft is
more than 6.8 m high and more than 4 m wide
Trajectory and Operations (Fig. 2). The complexity and mass of the space-
The Cassini-Huygens mission is a fruitful collab- craft is warranted by the ambitious program of
oration between ESA, NASA, and ASI which has scientific observations the spacecraft is
been investigating the Saturnian system since performing. Due to this and the power of the
2004, bringing spectacular new insights on the Titan IV Launcher, the spacecraft wasn’t injected
primary planet, its rings, and the exchanges within into a direct trajectory to Saturn but made use of
the system and within the Solar System. Particular gravity-assisted maneuvers at Venus, Earth, and
attention was devoted to the natural satellites, of Jupiter. These maneuvers increased the duration
which Titan, the largest moon, is a special target. of the voyage, which lasted about 7.5 years, but at
The mission has thus been instrumental in enhanc- the same time allowed to test the instruments
ing our understanding of the environment around during the different flybys, to improve their cal-
Saturn, in particular as to the habitability potential ibration, and also to make some interesting obser-
in bodies around giant planets. vations at Jupiter in conjunction with the
The Cassini-Huygens spacecraft consists of ▶ Galileo mission in orbit around the gas giant
two main elements: the NASA Cassini orbiter, in December 2000. This brief conjunction of the
named after the Italian-French astronomer two probes, complemented by simultaneous
Giovanni Domenico Cassini, who gave his observations from the Earth-orbiting Hubble
name to the ▶ Cassini Division in the rings and Space Telescope and Chandra X-ray Observa-
discovered several of Saturn’s major satellites, tory, provided an unprecedented opportunity for
and the ESA-provided Huygens probe, named the intimate study of the Solar System’s largest
after the Dutch astronomer, mathematician, and planet and enhanced our understanding of Jupi-
physicist Christian Huygens, who discovered ter’s radio emission and aurorae and their inter-
Titan in 1655. The Cassini-Huygens mission action with the solar wind. Jupiter’s
was launched on October 15, 1997, on a Titan magnetosphere and radiation belts and the inter-
IV-Centaur rocket from Cape Canaveral and actions between the magnetosphere and ▶ Io,
performed flybys of ▶ Venus, ▶ Earth, and ▶ Ganymede, and ▶ Europa were also explored.
▶ Jupiter before entering into orbit around Saturn The main body of the orbiter is nearly cylin-
on July 1, 2004 (Fig. 1). drical and consists of a lower equipment
Cassini-Huygens Space
Mission, Fig. 1 The Venus 1 flyby Saturn orbit
trajectory of the Cassini- 26 APR 1998 insertion
Huygens mission from Venus 2 flyby 1 JUL 2004
launch to Saturn orbit 24 JUN 1999
Saturn’s orbit
29.4 YEARS
insertion
11.8 YEARS
Jupiter’s orbit
Venus
targeting C
maneuver
SUN
3 DEC 1998
Launch
15 OCT 1997
Jupiter flyby
Earth flyby
30 DEC 2000
18 AUG 1999
Cassini-Huygens Space Cassini spacecraft

Mission, Fig. 2 Diagram 4m high-gain Low-gain
of the Cassini spacecraft antenna antenna (1 of 2)
carrying the Huygens probe
11m magnetometer
boom
Radar bay
Fields and
Radio/plasma wave particles pallet
subsystem antenna
(1 of 3) Huygens
Titan probe
Remote sensing
pallet
Radioisotope
thermoelectric
generator (1 of 3)
445 N engine (1 of 2)
platform, a propulsion module, and an upper charge-coupled device ▶ imaging system, a visi-
equipment platform, topped by a 4-m diameter ble/infrared mapping spectrometer, a composite
high-gain antenna. Attached about halfway up infrared spectrometer, a cosmic dust analyzer, a
the trunk is a remote sensing pallet, which hosts radio and plasma wave experiment, a plasma
cameras and other remote sensing instruments, spectrometer, an ultraviolet imaging spectro-
and a fields and particles pallet, which carries graph, a magnetospheric imaging instrument, a
instruments that study magnetic fields and magnetometer, and an ion/neutral mass spec-
charged particles. In order to point the instru- trometer. Telemetry from the communications
ments in the correct observing direction, the antenna and other special transmitters is also
entire spacecraft must be turned, although three used to make observations of the atmospheres of
of the instruments possess their own single-axis Titan and Saturn and to measure the gravity fields
articulation capability. of the planet and its satellites. On its orbit around
Cassini’s orbiter instrumentation (Fig. 2) con- Saturn, Cassini finds itself between 8.2 and 10.2
sists of a synthetic-aperture radar mapper, a astronomical units (AU) from the Earth. Because
of this, it takes between 68 and 84 min for radio

signals to travel from Earth to the spacecraft, and
vice versa.
More details and updates on the Cassini-
Huygens mission can be found at http://saturn.
jpl.nasa.gov/; http://sci.esa.int/cassini-huygens/;
http://www.nasa.gov/mission_pages/cassini/main/
Major Discoveries
The Cassini-Huygens mission has brought a host
of new and exciting discoveries in the Saturnian
system (Coustenis and Taylor 2008; Lorenz and
Mitton 2008; Brown et al. 2009; Dougherty
et al. 2009). They concern the distribution and
composition of the rings, the nature of the kronian
satellites and their interactions with the system,
the planet’s atmospheric envelope, and its
▶ magnetosphere. But also, the Cassini-Huygens Cassini-Huygens Space Mission, Fig. 3 This Cassini
discoveries have completely revolutionized our image shows a large hurricane-like storm at Saturn’s south
perception of the astrobiological and habitable pole. The dark eye of the “hurricane” spans about
8,000 km and it is surrounded by rings of clouds that
potential in a giant planet system, with strong
tower about 30–75 km above it. Contrary to the Earth,
implications also for exoplanetary worlds. Saturn’s storm is fixed in place – above the South
Given the long duration of the mission, the Pole – and is not powered by an ocean, since Saturn is a
complexity of the payload onboard the Cassini gaseous planet
Orbiter, and the amount of data gathered on the
primary planet, the satellites, and rings, it would clouds had not previously been seen on any planet
be impossible to describe all the new discoveries other than Earth. In addition, in counterpart, a
made (the reader can find detailed reviews in the strong six-sided jet stream was found at Saturn’s
books edited by Dougherty et al. 2009; Brown north pole known as “the hexagon” and
et al. 2009). We will therefore cite here only a few photographed again recently to span 32,000 km.
of the major breakthroughs showing how Saturn is probably best known for its system of
Cassini’s data have opened up a whole new chap- ▶ planetary rings, which makes it the most visu-
ter in Solar System exploration and in particular ally remarkable object in the solar system. They
have contributed to our understanding of the extend from 6,630 to 120,700 km above Saturn’s
astrobiological aspects of many of the Saturnian equator, average approximately 20 m in thick-
bodies (Lunine and Raulin 2010; Coustenis and ness, and are composed of 93 % ▶ water ice
Encrenaz 2013). with a smattering of ▶ tholin impurities and 7 %
In studying the primary planet, Cassini found ▶ amorphous carbon. The particles that make up
lightning on Saturn whose power is said to be the rings greatly vary in size. On September
approximately 1,000 times that of lightning on 20, 2006, a Cassini photograph revealed a previ-
Earth (Dougherty et al. 2009). In addition, in ously undiscovered planetary ring, outside the
October 2006, the probe detected an 8,000 km brighter main rings of Saturn and inside the
diameter hurricane with a six-sided eyewall at G and E rings. Apparently, the source of this
Saturn’s South Pole (Fig. 3). The storm’s eye is F ring is the result of the crashing of a meteoroid
about 2,000 km across with cloud speeds as fast off two of the moons of Saturn (Dougherty
as 530 km/h. Scientists believe that the storm is et al. 2009). Indeed, the interactions and material
the strongest of its kind ever seen. This observa- exchange between the rings and the satellites
tion is particularly notable because eyewall provide a significant breakthrough in our
Cassini-Huygens Space Mission, Fig. 4 Cassini cap- moon Prometheus periodically creates streamer-channels
tures the effects of the small moon Prometheus on two of in the F ring, as can be seen on the left of the image.
Saturn’s rings in this image taken on July 30, 2009, at a Prometheus is overexposed in this image. Bright specks in
distance of approximately 1.8 million kilometers from the image are background stars. This view looks toward
Saturn. A long, thin shadow cast by the moon stretches the northern, dark side of the rings from about 28 above
across the A ring on the right. The gravity of the small the ring plane
understanding of the Saturnian system provided the main Cassini division, other gaps have been
by the Cassini spacecraft (Fig. 4). Thus, on Octo- found in the rings where particle density drops
ber 6, 2009, another discovery was announced of significantly. A couple of them have identified
a tenuous outer disk of material in the plane of origins by known moons embedded within
▶ Phoebe’s orbit. The ring is from 128 to them, while others are at locations of known
207 times the radius of Saturn and is thought to ▶ orbital resonances with ▶ Saturn’s other
originate from micrometeoroid impacts on the moons. Finally, still some gaps remain
satellite Phoebe, which orbits at an average dis- unexplained to date.
tance of 215 Saturn radii. The ring material So far, the mission has discovered and con-
should thus share Phoebe’s retrograde orbital firmed several new satellites, bringing the number
motion and after migrating inward would up to 62. In the satellite system, the Cassini orbiter
encounter ▶ Iapetus’ leading face, which could has produced a large range of new results, demon-
help explain the dramatic two-faced nature of this strating that these small bodies are far from being
satellite. While the infalling material cannot be icy, dead worlds but rather active bodies with
directly responsible for the observed pattern of resurfacing and cryovolcanic features. The
light and dark regions on Iapetus, it is believed to moons of Saturn are a diverse collection (Fig. 5).
have initiated a runaway thermal self-segregation Cassini has explored their icy landscapes in
process in which ice sublimes from warmer unprecedented detail, solving long-standing mys-
regions and condenses onto cooler regions. This teries and sharing many new surprises: ▶ Iapetus
leaves contrasting areas of dark ice-depleted res- has an enormous ridge along its equator, in addi-
idue and bright ice deposits. Another important tion to its two sides of remarkably different bright-
discovery was that the source of the material in ness. ▶ Rhea may have its own faint rings. And
the E ring, the second outermost and very wide, sponge-looking Hyperion is so porous that impacts
was probably the water-laden plumes emanating tend to just punch into the surface, and its gravity
from the “tiger stripes” of the south polar region is so low that what material does get ejected tends
of the cryovolcanic satellite Enceladus. Besides to leave the moon altogether.
Cassini-Huygens Space Mission, Fig. 5 On June lower right. The view was obtained at a distance of
28, 2007, the Cassini cameras captured this trio of icy approximately 291,000 km from Enceladus, looking
moons against Saturn’s atmosphere and rings toward the sunlit side of the rings from about a degree
(Dougherty et al. 2009). Enceladus is located on the below the ringplane. Scale in the image ranges from 17 km
planet’s shadow-draped limb at the center; Pandora is a per pixel on Enceladus to 32 km per pixel on Saturn in the
bright speck hovering near the rings; and Mimas is seen at background
A big surprise came from February 2010 in several of the instruments’ data, not only on
observations of ▶ Mimas, the satellite character- Titan and ▶ Enceladus, but also on Iapetus and
ized by the 130-km wide Herschel crater domi- Phoebe and ▶ Dione, with Cassini VIMS show-
nating its surface, showing large temperature ing that the dark material in the Saturn system
differences across the surface with no surface could be organics (Dougherty et al. 2009).
brightness features to explain it. The differences More surprises came from the combined
are believed to be due to Mimas’ thermal inertia investigations of the orbiter and of the Huygens
being considerably higher in the colder regions probe in the case of Titan as shown for example in
than the hotter. In such a case, heat could soak Fig. 6 for the Huygens landing site (Coustenis
into the Mimas interior more easily, rather than and Taylor 2008; Lorenz and Mitton 2008;
raise the temperature of the surface. This sup- Mueller-Wodarg et al. 2014) but also from very
poses that the thermal conductivity of the satellite close Cassini flybys of Enceladus (Brown
has to be at least ten times greater in the cold et al. 2009). In some ways, the moons Titan and
regions with respect to the warmer regions, while Enceladus have turned out to be unique revela-
there are no apparent differences in the visible- tions of the Cassini mission, making the Satur-
wavelength observations. These facts and the nian system’s exploration very relevant to the
giant crater, Herschel, residing within this cold search for ▶ life and habitable conditions in the
region remain a mystery. Solar System. Titan, with its thick organic-laden
Cassini also discovered the presence of atmosphere, clouds, dunes, and rivers and lakes
organics (hydrocarbons mainly) on several of of liquid ▶ methane-ethane on its surface (Stofan
Saturn’s satellites seen through their signatures et al. 2007; Brown et al. 2008; Raulin et al. 2012),
Mission, Fig. 6 Artwork
showing the Cassini orbiter
observing the Huygens
probe landing site with
VIMS and UVIS in 2008
is a rich laboratory for chemistry and processes occurred on the prebiotic Earth. No other place
that may resemble early Earth but with different in the solar system has this type of ongoing
materials in play (Coustenis and Taylor 2008). chemistry. The Cassini-Huygens era of investi-
Furthermore, the presence of an internal liquid gation has furthered our understanding of Titan as
water ocean is more substantiated with new mea- the largest ▶ abiotic organic factory in the Solar
surements. And with its towering south polar System.
plume of icy particles and water vapor, Enceladus Some essential breakthroughs related to
is a geologically active moon, certainly a strong Astrobiology within the realm of Saturn are sum-
candidate for liquid water underneath its surface marized hereafter and on Fig. 7.
and energy sources, and incredibly important for
the study of potentially habitable environments. Titan’s Atmosphere
Both of these moons are compelling targets for With the current picture of Titan’s organic chem-
future exploration in that context, but in the istry, the chemical evolution of the main atmo-
meantime, Cassini-Huygens is providing us with spheric constituents – ▶ dinitrogen (N2) and
major insights bearing high relevance to Astrobi- ▶ methane – produces complex refractory
ology, the ▶ origin of life, and habitability. organics through ▶ photolysis and ▶ photochem-
istry. Measurements throughout the Titan atmo-
Titan sphere, both remotely and in situ, have indicated
The Cassini orbiter and the Huygens descent the presence of numerous hydrocarbon and
probe were designed to be part of a common ▶ nitrile gases, as well as a complex layering of
strategy to uncover the mysteries shrouding organic ▶ aerosols that persists all the way down
the enigmatic satellite of Saturn, ▶ Titan. Titan to the surface of the moon. The organic chemistry
is an organic paradise that is certain to tell us detected in the higher atmosphere by the Ion and
much about the chemical evolution that may Neutral Mass Spectrometer (INMS, Waite
lead to life. Water ice and ▶ carbon dioxide ice et al. 2007) provided feedback and useful infor-
have been reported to exist currently on the sur- mation for all studies and models of the satellite’s
face. Transient episodes of melting of the water chemical composition, complemented by mea-
ice by either geologic activity or impacts would surements in the ▶ stratosphere made by the
expose organics to aqueous alteration, as well as Composite Infrared Spectrometer (CIRS,
contact with carbon dioxide, leading potentially Coustenis et al. 2010) and by the chemistry infer-
to reaction pathways that mimic those that ences from the Gas Chromatograph-Mass
Mission, Fig. 7 Some of
the major discoveries that
the Cassini-Huygens
mission made on Titan:
transient atmospheric
phenomena (clouds) and
several geomorphological
features (dunes,
cryovolcanoes, channels,
mountains and lakes) are
shown in this composite
(Credit: G. Tobie)
Spectrometer (GC-MS, Niemann et al. 2010), as composition have been recorded by the Cassini
well as density and temperature data retrieved by instruments.
the Huygens Atmospheric Structure Instrument The products accumulate on the surface,
(HASI, Fulchignoni et al. 2005) during the together with condensed volatile organic com-
descent to the surface near Titan’s equator. The pounds such as HCN and benzene. The abun-
highly complex organic species in the ionosphere dance of methane and its organic products in the
found by INMS are the precursors of the hydro- atmosphere, seas, and dunes exceeds by more
carbons and nitriles found in the stratosphere, than one order of magnitude the carbon inventory
which form aggregates and eventually condense in the Earth’s ocean, biosphere, and fossil fuel
out on the surface. Some of these chemical com- reservoirs. Indeed, the measured value of the
ponents are molecules of prebiotic interest (like irreversible conversion of the methane in the
hydrogen cyanide and HCN, Raulin et al. 2008). atmosphere into higher-order organic/nitrile
Thus, it appears that Titan is a chemical factory in compounds that eventually end up deposited on
which the formation of complex positive and the surface of Titan is near that of our terrestrial
negative ions is initiated in the high thermosphere reference, indicating that methane is resupplied
as a consequence of magnetospheric- and converted at a rate that prevents the buildup
ionospheric-atmospheric interactions involving of the heavier isotopologue over time as is the
solar EUV, ▶ UV radiation, and energetic ions case of ▶ nitrogen. However, the source of the
and electrons. Furthermore, seasonal variations atmospheric methane remains a mystery today
of the atmospheric temperature and chemical although various models and hypotheses have
been advanced, including its periodic outgassing and exciting terrain. One of the most efficient
from the interior via cryovolcanism. applications of the synergy between the orbiter
The Cassini cameras and spectrometers, as and the probe is the mapping of Titan’s surface.
well as the Huygens instruments, showed the While the Cassini orbiter provided detailed views
presence of clouds and storms in Titan’s atmo- of Titan’s surface with its camera, mapping spec-
sphere and described the distribution of the aero- trometer, and radar, the Huygens probe,
sols (haze) throughout the atmosphere. Most of descending through the atmosphere on January C
the clouds detected in the past years were in 14, 2005, returned extraordinarily detailed
Titan’s southern hemisphere, as expected given images with resolutions ranging from 10 m at
the season on Titan that has been essentially 10 km down to centimeters at the surface
probed (summer in the south), which means that (Fig. 7). The Huygens Atmospheric Structure
solar heating is concentrated there. In recent Instrument (HASI) gave the conditions of pres-
years, however, they have been found to be build- sure, density, and temperature throughout Titan’s
ing up also in the north as the season changes. atmosphere and in particular on Titan’s surface to
Furthermore, temporal and even seasonal varia- be 1.5 bar and 93.7 K (Fulchignoni et al. 2005),
tions in the thermal structure and the chemical serving as a reference for many studies.
composition of Titan have been recorded by the For the surface discoveries, the context is pro-
Cassini instruments. During the Cassini mission, vided by the Cassini radar (RADAR), the Visual
Titan’s season moved from northern winter to and Infrared Mapping Spectrometer (VIMS), and
northern spring, returning to the same configura- the Imaging Science Subsystem (ISS) data (Porco
tion as during the Voyager encounter sometime in et al. 2005), while the ground truth was obtained
2010. The same enhancement in the northern by several of the Huygens instruments at the
polar composition was found for most gases, but probe’s ▶ landing site, like the images and spec-
since mid 2012, a reversal has been monitored as tra of Descent Imager/Spectral Radiometer
the south pole is now coming into winter (DISR, Tomasko et al. 2005) or the composition
exhibiting stronger signatures of gases and haze. measurements of the ▶ GC-MS (Niemann
The seasonal variations of the haze, whose verti- et al. 2010). Radar observations suggest that the
cal structure has changed with the detached layer ultimate fate of this aerosol precipitation is the
dropping in altitude significantly in recent years, generation of expansive organic-laden dunes
can be attributed to a varying Hadley circulation. (Lorenz et al. 2006; Radebaugh et al. 2008) that
Another important discovery was that the strato- were observed around Titan’s equator for the first
spheric zonal winds and temperatures in both time by Cassini (Fig. 7). These dunes are remark-
hemispheres of Titan are symmetric about a able in being exactly the same size and shape as
pole that is offset from the surface pole by ~4 . linear (longitudinal) dunes on Earth such as those
Cassini-Huygens has also provided important found in the Namibian and Saharan desert and are
information on the origin and evolution of Titan’s essentially found near Titan’s equator. Indeed,
atmosphere by measuring the ▶ noble gas con- the mid latitude regions around the equator on
centrations (like argon for the first time) and their Titan were found to be rather uniformly dark with
isotopic abundances, as well as the nitrogen and some extensive bright regions (mostly elevated
carbon stable isotopic ratios. These measure- structures like mountains and ridges), while the
ments also provide important clues about the poles are relatively bright but also filled with
overall role of escape, chemical conversion, large dark areas now recognized as hydrocarbon
outgassing, and recycling in the evolution of “lakes” (mostly in the north pole) formed possi-
Titan’s atmosphere. bly by precipitation through the atmosphere. Few
craters were detected on Titan’s surface, which
Titan’s Surface then appears rather young.
Only with Cassini-Huygens did it become possi- Radar-bright channels (probably cobbled
ble to acquire a clear picture of Titan’s complex streambeds like that at the Huygens landing
site) have been observed at low and In 2013, a study even reported the possible pres-
mid-latitudes, while channels incised to depths ence of blocks of hydrocarbon ice floating on the
of several hundred meters are seen elsewhere, surface of these lakes.
and at high latitudes, radar-dark, meandering Thus, the diversity of the terrains on Titan
channels are seen that suggest a lower-energy depicted by the Cassini-Huygens instruments
environment where deposition of fine-grained includes a host of geologic features (Fig. 7):
sediment occurs (Soderblom et al. 2007a; Lorenz
et al. 2008a). Fluvial modification of the surface • Erosional features such as channels and den-
was very evident at the Huygens landing site dritic networks, possible lakes and seas, fluvial
(Tomasko et al. 2005). Radar and near-infrared erosional deltas and other erosional and depo-
imagery has revealed channels on much larger sitional constructs such as dunes (Radebaugh
scales than those seen by Huygens. et al. 2008), possible glacial-flow
Furthermore, in July 2006, Cassini found the constructs, etc.
first proof of hydrocarbon lakes near Titan’s • Impacts: the very low crater frequency is
north pole (Stofan et al. 2007), which was con- indicative of active geological surface
firmed in January 2007 (Mitri et al. 2007; Hayes processes
et al. 2008). In March 2007, additional images • Volcanotectonic features: domes, possible
near Titan’s north pole discovered hydrocarbon cryovolcanic flows, and bright spots (Sotin
“seas.” These very dark features at the high north- et al. 2005; Nelson et al. 2007; Lopes
ern latitudes of Titan were finally shown to be et al. 2013) as well as mountain chains
liquid-filled (most probably with ethane-rich (Radebaugh et al. 2007), many of these fea-
mixtures, Brown et al. 2008; Raulin 2008, 2012; tures may be active regions on Titan’s surface
Cordier et al. 2012) basins – “lakes.” The features
range in size from less than 10 km2 to at least The features Cassini-Huygens discovered on
100,000 km2. They are confined to the region Titan’s surface were more complex than any
poleward of 55 N. To date, some 655 such fea- expected, with landforms that seem to resemble
tures have been identified and mapped (Fig. 7). the landscapes on Earth, including hills, dunes, a
Other small lakes exist more to the south, like the deflated lakebed, but all composed of completely
Ontario Lacus. In 2010, specular reflections of different constituents, many of which could be
the Sun off of Titan’s northern lakes (“sunglints”) ices and organic material. The ambient condi-
were reported, while Cassini looked back toward tions and direct measurement of methane evapo-
a crescent Titan at high phase angle. The data rating from under the landed probe imply that the
were used to set some constraints on possible working erosive agent is liquid methane, not liq-
surface waves on a north polar lake (Jingpo uid water.
Lacus), and since then observed specular reflec- The surface of Titan, as revealed by the
tions have been quite useful in corroborating the Cassini orbiter and the Huygens probe, offers us
presence of large extents of liquid hydrocarbons an opportunity to stretch our current models in an
in Titan’s surface. The major seas on Titan, effort to explain the presence of dunes, rivers,
which are very extended in size, are namely Kra- lakes, cryovolcanoes, and mountains in a world
ken Mare (surface area of at least 400,000 km2 where the rocks are composed of water ice rather
centered at 68 N, 310 W), Ligeia Mare (79 N, than silicates and the liquid is methane or ethane
248 W), and Punga (85 N, 339 W). Although rather than liquid water.
some of these “seas” or “lakes” or even “oases” Titan’s tectonism involves a number of very-
are found at mid latitudes, most are located in the large-scale linear features seen optically, notably
northern hemisphere at Titan’s current season, the dark dune-filled basins. Some linear mountain
but they are expected according to some models ranges have been detected, several forming a
to build up in the south, following the atmo- chevron pattern near the equator, with a large
spheric evolution, as the season changes. bright terrain (Xanadu) extending over
Mission, Fig. 8 Cassini’s
major breakthroughs in our
understanding of the
Enceladus environment are
connected to the southern
polar region, where the
magnetometer and the C
cameras detected plumes of
gas and ice particles
emanating from a series of
warm fractures, dubbed the
“tiger stripes”
3,400 km in diameter. RADAR/SAR (synthetic- astrobiological potential of liquid water erupting

aperture radar (SAR)) imagery shows Xanadu to onto photochemically produced organic mole-
be extremely rugged. cules. Several likely cryovolcanic structures
As said before, the N2-CH4 by-products in have been identified in Cassini near-infrared and
Titan’s atmosphere eventually end up as sedi- radar images (Fig. 8). Although definitive evi-
ments on the surface, where they accumulate dence for active volcanism has not yet been pro-
presently at a rate of roughly 0.5 km in 4.5 Ga. duced, there are apparent surface changes in
Since no large source was detected by Cassini to Cassini data that require explanation.
resupply methane, cryovolcanic outgassing has Titan’s overall density (1.88 g/cm3) requires it
been hypothesized, yet over what timescales and to have roughly equal proportions of rock and ice.
through which internal processes is unknown. After its accretion, Titan was probably warm
Cassini-Huygens also found that the balance of enough to allow differentiation into a rocky core
geologic processes – impacts, tectonics, fluvial, with a water/ice envelope, but whether an iron or
aeolian – is somewhat similar to the Earth’s, more iron-sulfur core formed during the subsequent
so than for Venus or Mars. Titan may well be the evolution remains uncertain. In addition, the
best analogue to an active terrestrial planet in the presence of an internal liquid water ocean for
sense of our home planet, albeit with different Titan is supported by models, radar and gravity
working materials (Solomonidou et al. 2013; Cassini measurements, and the HASI experi-
Mueller-Wodarg et al. 2014). ment. The extremely low-frequency electric sig-
In addition, the detection of Argon 40, and nal recorded by HASI was interpreted as a
observations of what appear to be flows from Schumann resonance between the ionosphere
cryovolcanoes, suggests that the interior of and a modestly conducting ocean roughly 50 km
Titan is geologically active; theoretical calcula- below the surface. Thermal evolution models
tions suggest a heat flow at present of about 8 % suggest that Titan may have an icy crust between
that of the Earth, sufficient to mobilize water as 50 and 150 km thick, lying atop a liquid-water
liquid in the interior as the working fluid for ocean a couple of hundred kilometers deep, with
▶ cryovolcanism. Cryovolcanism is a process of some amount (a few to 30 %, most likely 10 %)
particular interest at Titan because of the known of ammonia dissolved in it, acting as an
Cassini-Huygens Space Mission, Fig. 9 One of the Cassini flybys of Enceladus when the spacecraft flew at quite low
altitudes into the satellite’s plumes (artistic impression)
antifreeze. Beneath lies a layer of high-pressure a cloud around the moon and feeding the adjacent
ice. Cassini’s measurement of a small but signif- E ring. The Cassini spacecraft provided definitive
icant nonsynchronous contribution to Titan’s proof that Enceladus is currently geologically
rotation is most straightforwardly interpreted as active when multiple Cassini instruments
a result of decoupling of the crust from the deeper detected plumes of gas and ice particles emanat-
interior by a liquid layer (Lorenz et al. 2008b). ing from a series of warm fractures centered on
The presence of ammonia, from which Titan’s the south pole, dubbed the “tiger stripes” (Fig. 8).
nitrogen atmosphere was presumably derived, Cassini data strongly suggested the presence
distinguishes Titan’s thermal evolution from of liquid-water reservoirs that erupt in geysers on
that of Ganymede and Callisto. The recent detec- Saturn’s moon Enceladus. Images had also
tion of periodic tidal stresses on Titan requires shown particles of water in its liquid state emitted
that its interior be deformable over timescales of by icy jets and towering plumes. Several flybys of
the orbital period in a way that is consistent with a Enceladus followed bringing more data and
global ocean at depth (Iess et al. 2012). All of revealing an extremely intriguing world (Fig. 9).
these indications make Titan a very likely habit- In March 2008, Cassini swept by Enceladus’
able environment. south pole at an altitude of 52 km and plunged
into a south polar plume, scooping up particles
Enceladus and gases to sample their composition. The gases
When Cassini had its first encounter with that were “tasted” by Cassini’s INMS bore a
▶ Enceladus on February 17, 2005, the magne- strong resemblance to the gases that issued from
tometer instrument saw a bending of Saturn’s comets. All of the “fields and particles” Cassini
magnetic field, with the plasma being slowed instruments (the ones that measure the abun-
and deflected as it passed Enceladus. Data col- dance, compositions, and motions of plasma,
lected during the March 9, 2005 flyby provided ions, atoms, molecules, particles, and magnetic
further evidence of the existence of an atmo- fields in situ, wherever Cassini travels) have been
sphere around the southern pole of Enceladus. sampling the plumes (Fig. 9).
The cosmic dust analyzer recorded thousands of Enceladus is thus the second cryovolcanically
hits from tiny particles of dust or ice coming from active icy satellite that has been identified (Triton
is the only other known active icy satellite, but The two major themes in Titan exploration – the
the process driving its cryovolcanism may not be methane cycle as an analogue to the terrestrial
linked to an internal heat source) and can be used hydrological cycle (Atreya et al. 2006) and the
to study active processes that are thought to have chemical transformations of ▶ complex organic
once played an important role in shaping the molecules in the atmosphere and the
surfaces of other icy satellites. These processes surface – render Titan a very high priority if we
include tidal heating, cryovolcanism, and ice tec- are to understand how volatile-rich worlds evolve C
tonism, which all can be studied as they currently and how organic chemistry and planetary evolu-
happen on Enceladus. Moreover, the plume tion interact on large spatial and temporal scales.
source region on Enceladus samples a warm, Both are of keen interest to planetology and
chemically rich, environment that may facilitate astrobiology.
complex organic chemistry and biological pro- To answer the several remaining vital ques-
cesses. CIRS on Cassini has demonstrated that tions that Cassini has raised for Titan and for
the Enceladus south pole was the warmest portion Enceladus, a new mission (the Titan Saturn Sys-
of this moon, considerably much warmer than the tem Mission, TSSM) was proposed and studied in
equator (Spencer 2013). 2008 by both ESA and NASA. This ambitious
Enceladus is arguably a place in the solar mission could bring the required long-term
system where exploration is most likely to find a exploring capabilities combing an orbiter and
demonstrably habitable environment, and several two in situ elements (a montgolfière balloon and
researches point to the possibility that Enceladus’ a lander) with state-of-the-art technology and
plumes, tectonic processes, and possibly a liquid- instruments (see www.lesia.cosmicvision/tssm/
water reservoir beneath the surface may create a tssm-public). After this study, other, simpler but
complete and sustainable geochemical cycle that also exciting, mission concepts have also been
may allow it to support life. While other moons in proposed for a return to the Saturnian system
the solar system have liquid-water oceans cov- within the next two or three decades to study
ered by kilometers of icy crust, in the case of Titan with orbiters, lake landers, balloons,
Enceladus, the pockets of liquid water may be airplanes, etc.
no more than tens of meters below the surface.
Cassini has thus discovered a new potential
▶ habitat in our Solar System, well outside the See Also
traditional ▶ habitable zone, which needs to be
extended to include these “deep habitats.” ▶ Aerosols
▶ Cassini
▶ Cassini Division
Future Directions ▶ Complex Organic Molecules
▶ Enceladus
The primary mission for Cassini ended on July ▶ Galileo Galilei
30, 2008. However, given the excellent condition ▶ Giant Planets
of the orbiter, the mission was extended to 2010. ▶ Habitable Zone
On February 3, 2010, NASA announced another ▶ Hubble Space Telescope
extension for Cassini, this one for 6–1/2 years ▶ Huygens
until 2017. The extension enables another ▶ Huygens Probe
155 revolutions around the planet, 54 flybys of ▶ Iapetus
Titan, and 11 flybys of Enceladus. ▶ Isotopic Ratio
However, even after these extensions, several ▶ Landing Site
questions and scientific themes remain that can- ▶ Magnetosphere
not be addressed by Cassini in its current config- ▶ Methane
uration or with its present instrumentation. ▶ Mimas
▶ Organic Material Inventory Lorenz RD, Mitton J (2008) Titan unveiled. Cambridge
▶ Origin of Life University Press, Cambridge
Lorenz RD et al (2006) The sand seas of Titan: Cassini
▶ Photochemistry, Atmospheric RADAR observations of longitudinal dunes. Science
▶ Photolysis 312:724–727
▶ Planetary Rings Lorenz RD et al (2008a) Fluvial channels on Titan: initial
▶ Rhea Cassini RADAR observations. Planet Space Sci
56:1132–1144
▶ Satellite or Moon Lorenz RD et al (2008b) Titan’s rotation reveals an inter-
▶ Saturn nal ocean and changing zonal winds. Science
▶ Terrestrial Analog 319:1649–1651
▶ Tholins Lunine J, Raulin F (2010) Titan and the Cassini-Huygens
mission, Chapter 29. In: Gargaud M, Lopez-Garcia P,
▶ Titan Martin H (eds) Origin and evolution of life: an astro-
▶ Volatile biology perspective. Cambridge University Press,
New York
Mitri G, Showman AP, Lunine JI, Lorenz RD
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Titan’s upper atmosphere. Science 316:870–875
Catabolism is the subset of metabolic networks
by which organic compounds are degraded to
simpler organic or inorganic compounds. Catab-
C-Asteroid olism includes oxidation reactions – coupled to
the reduction of coenzymes such as NAD+ or
Alan W. Harris FAD – and sometimes substrate-level phosphor-
DLR, Institute of Planetary Research, Berlin, ylation steps leading to ATP synthesis.
Germany
See Also
Definition
▶ Anabolism
▶ Asteroid taxonomic systems are based on spec- ▶ Metabolism
tral features observed in reflected sunlight, with
letters of the alphabet used to denote different
taxonomic types. A C-type asteroid is one with
a relatively flat and featureless reflection spec- Cataclysmic Pole Shift Hypothesis
trum in visible light, similar to that of ▶ carbona-
ceous chondrite ▶ meteorites and with a low ▶ True Polar Wander, Theory of
▶ albedo (typically 0.03–0.1), characteristics
indicative of a carbonaceous (carbon-rich) min-
eralogy. C-type asteroids are common in the
outer main belt but are also present in the near- Catalyst
Earth asteroid population. Many C-type asteroids
have weak absorption features in the near- Henderson James (Jim) Cleaves II
infrared region of the spectrum, apparently due Earth–Life Science Institute (ELSI), Tokyo
to the presence of water-bearing ▶ minerals. Institute of Technology, Meguro–ku, Tokyo,
Japan
See Also Blue Marble Space Institute of Science,
Washington, DC, USA
▶ Albedo Center for Chemical Evolution, Georgia Institute
▶ Asteroid of Technology, Atlanta, GA, USA
▶ Carbonaceous Chondrite Definition
▶ Meteorites
▶ Mineral In chemistry, catalysts are compounds that
▶ Near-Earth Objects increase the rate of a reaction but are not
398 Catena, Catenae
consumed in the reaction, as a reagent would See Also

be. They can therefore generally participate in
reactions multiple times. The term “catalyst” is ▶ Clay
derived from the Greek katalύein, meaning to ▶ Enzyme
dissolve or loosen. Catalysts may be metals, min- ▶ Protein
eral surfaces, ▶ enzymes, ▶ ribozymes, or small ▶ Ribozyme
organic molecules, among other possibilities.
Overview
Catena, Catenae
Catalysts operate by changing the rate-limiting
free energy change to the transition state relative Roland J. Wagner
to that of the corresponding uncatalyzed German Aerospace Center (DLR), Institute of
reaction, resulting in a larger reaction rate at a Planetary Research, Berlin, Germany
given temperature. The physical mechanism of
catalysis can be complex. Catalysts may affect
the reaction environment or bind to the reagents Synonyms
to polarize bonds forming intermediates that
do not occur in the cognate uncatalyzed reac- Crater chain
tions. Catalysts have no effect on the chemical
equilibrium of a reaction because the rates of
both the forward and reverse reactions are
affected. Definition
In catalyzed reactions, as in uncatalyzed reac-
tions, the overall reaction rate depends on the A catena is a linear, slightly curved, or sinuous
frequency of collision of the reactants in the chain of circular to elliptical depressions. The
rate-limiting step. The catalyst usually acts in depressions can be surrounded by raised rims
this limiting step, and rate changes are propor- and are contiguous or separate. A crater chain
tional to the amount of catalyst present. As cata- may originate from volcanic or impact processes.
lysts are not consumed in a reaction, only small Depressions in volcanic catenae are generally
amounts may be needed to increase the rate of the rimless. Raised rims and a more-or-less uniform
reaction significantly. In reality, however, cata- size distribution of the depressions are character-
lysts are sometimes inhibited, deactivated, or istic of impact crater chains. Impact catenae are
destroyed via secondary processes. Substances created (a) either by fragments of a projectile that
that reduce the activity of catalysts are called disintegrated prior to impact or (b) by material
inhibitors if the reduction is reversible and poi- ejected when a crater is formed, forming chains
sons if the reduction is irreversible. of secondary craters.
Catalysts can be heterogeneous or homoge-
neous, depending on whether they exist in the
same phase as the substrate. Heterogeneous cat-
alysts act in a different phase than the reactants, See Also
for example, solids that act on liquid or gaseous
substrates. Homogeneous catalysts function in ▶ Crater, Impact
the same phase as the reactants, for example, ▶ Fumarole
cytosolic enzymes are examples of homogeneous ▶ Olympus Mons
catalysts. ▶ Patera, Paterae
CCD 399
Definition
Cavitation Zone
CCD is the abbreviation for charge-coupled
▶ Spallation Zone device. A CCD is a photoelectronic imaging
device commonly used for astronomical observa-
tions in the visible domain. A CCD is a solid-state
silicon-based detector where each of the numer- C
Cavus, Cavi ous pixels (up to ten million) stores in a potential
well the electrons produced by photons (one to
Ernst Hauber one). At the end of the exposure, charges are
Deutsches Zentrum f€ur Luft- und Raumfahrt transferred from one pixel to the next, up to the
(DLR) e.V., Institut f€ur Planetenforschung, output amplifier, by manipulation of voltages
Berlin, Germany applied to surface electrodes (see Fig. 1).
Back-illuminated (or thinned) CCDs feature an
Definition
Electrodes
A cavus is a hollow, irregular steep-sided depres-
sion occurring usually in arrays or clusters SiO2
(definition by the International Astronomical
Union; http://planetarynames.wr.usgs.gov/jsp/
append5.jsp). It is used as a descriptor term for
naming surface features on ▶ Mars.
Substrate (doped Silicon)
See Also
SiO2
▶ Mars
CC
SiO2
CCD
Daniel Rouan
CCD, Fig. 1 The principle of charge transfer from one
pixel to the next one in a CCD. Top: photons penetrate the
Meudon, France substrate and create electrons. Middle: the electrons are
attracted and stored below an electrode polarized with a
positive voltage and isolated from the substrate, thanks to
a thin layer of silicon oxide. Bottom: a voltage is applied to
Synonyms the next electrode and reset on the first one so that elec-
trons travel one step. The procedure is repeated until all
Charge-coupled device pixels are read at the end of the chain
400 CD
excellent quantum efficiency for wavelengths up

to l = 1 mm. Because of their better efficiency Cell
and the convenience of obtaining directly a digi-
tal image at the output, CCDs definitely replaced Angeles Aguilera
photographic plates for use in astronomy in the Laboratorio de Extremófilos, Centro de
1980s, especially since it has been possible to tile Astrobiologı́a (INTA-CSIC), Torrejón de Ardoz,
side by side several CCDs (up to several tens) and Madrid, Spain
produce cameras with a wide sensitive surface.
Keywords
See Also
Cell structure; Endosymbiotic theory; Micros-
▶ Imaging copy; Organelles
Definition
CD The cell is the smallest unit of living matter

capable of performing all the activities necessary
▶ Circular Dichroism for life (Alberts et al. 1994). In fact it is the
smallest structure with a complete metabolism
because it has all the physical and chemical com-
ponents needed for its own maintenance and
Celestial Equator growth. All living organisms are made of cells.
The simplest forms of life are individual cells that
Daniel Rouan propagate by division, while more complex
LESIA, Observatoire Paris-Site de Meudon, organisms are multicellular, that is, their bodies
Meudon, France are cooperatives of many kinds of specialized
cells that could not survive for long time by
themselves.
Definition
The celestial equator is the projection of the History

Earth’s equator on the sky, represented as a
great circle on the imaginary celestial sphere. The first person to use the word “cell” was Robert
As a result of the Earth’s axial tilt, the celestial Hook (1665) who described what he called the
equator is inclined by 23.5 with respect to the cella in a piece of cork (Fig. 1). He used this
ecliptic plane. The celestial equator is the origin term because the cork appeared to be composed
of ▶ declination, one of the two coordinates used of thousands of small chambers that resembled the
to locate an object on the sky. individual sleeping rooms in monasteries. How-
ever, Antonie Philips van Leeuwenhoek
(1632–1723) using his handcrafted microscopes
See Also was the first to observe and describe single-celled
microbial organisms, which he originally referred
▶ Coordinate Systems to as animalcules. After Van Leeuwenhoek, in
▶ Declination 1838 a botanist, Matthias Jakob Schleiden, and a
▶ Ecliptic zoologist, Theodor Schwann, formally proposed
Cell 401
Cell, Fig. 1 Portraits of

Robert Hook and Antonie
Philips van Leeuwenhoek,
two of the founders of cell
biology and microbiology.
Below them, the portraits of
Matthias Jakob Schleiden
and Lynn Margulis, known C
for their cell theory and
endosymbiotic theory
respectively
“The Cell Theory” stating that: (1) all organisms Overview

are composed of one or more cells, (2) all cells
come from preexisting cells, (3) vital functions of At first sight, cells exhibit a staggering diversity.
an organism occur within cells, and (4) all cells Some lead a solitary existence, others live in
contain the hereditary information necessary for communities; some have defined geometric
regulating cell functions and for transmitting shapes, others have flexible boundaries; some
information to the next generation of cells. Their swim, some are sedentary. However, all cells
theory, which nowadays seems so obvious, was a have several basic features in common: they are
milestone in the development of modern biology. bounded by a plasma membrane that physically
The cell theory was extended in 1855 by Rudolf separates them from the outside environment.
Virchow, who stated that new cells come into Within the membrane is a semifluid substance,
existence only by the division of previously cytosol, composed mostly of water (70–90 %). In
existing cells. Cells cannot arise by spontaneous the cytosol is where a variety of specialized struc-
generation from nonliving matter (Campbell and tures named ▶ organelles are located. All cells
Reece 2002). contain also chromosomes, carrying genes for
402 Cell
Cell, Fig. 2 Although all the cells have several basic The cells are cylindrical with a rounded anterior and
features in common, there is a great variety of specific tapered posterior. The chloroplasts are well-developed
cellular body plans. Some cells are organisms in their own and can glide and swim using their flagella. (e) White
right, and some make up the bodies of multicellular organ- and red blood cells also referred to them as leucocytes
isms. (a) Mixed diatoms, these algae are single-cell or and erythrocytes principal cells of the immune system
colonial and are surrounded by a rigid siliceous envelope involved in defending the body against both infectious
named theca. (b) Bacterial cells belonging to the species disease and foreign materials and cells involved in deliv-
Escherichia coli a rod-shaped bacterium that is commonly ering oxygen to the body tissues via the blood flow
found in the lower intestine of warm-blooded organisms. through the circulatory system. (f) Spyrogyra cells, a
(c) Amoeba cell, this small protozoan uses tentacular pro- filamentous green algae named for the helical or spiral
tuberances called pseudopodia to move and phagocytose arrangement of the chloroplasts. It is commonly found in
smaller unicellular organisms, which are enveloped inside freshwater areas, and there are more than 400 species of
the cell’s cytoplasm in a food vacuole, where they are Spirogyra in the world. Spirogyra measures approxi-
slowly broken down by enzymes. (d) Euglena cell, a mately 10–100 mm in width and may stretch
photosynthetic protist with at least 150 described species. centimeters long
synthesizing all the proteins needed for cell simplest, and the more complex eukaryotic cell
growth, repair, and reproduction. (Fig. 2). By definition, prokaryotes are those
Despite all the different cell morphologies, it organisms whose cells are not subdivided by
is surprising that there are only two types of cells. membranes into a separate nucleus and cyto-
Based on differences in compartmentalization, plasm. All prokaryote cell components are
cells can be divided into prokaryotic cell, the located together in the same compartment; the
Cell 403
genetic material (DNA) is concentrated in a few as 500. A typical prokaryotic cell is rod
region called the nucleid, but no membrane sep- shaped and about 2–3 mm long and 1 mm wide
arates this region from the rest of the cell. On the (1 mm = 0.001 mm). However, bacteria are not
contrary, eukaryotic cells contain a membrane- limited to a rod shape; spherical, filamentous, or
bound organelle named nucleus where the spirally twisted bacteria are also found. Eukary-
genetic material is contained. Only bacteria and otic cells are generally much bigger than pro-
archaea are prokaryotic cells. Protists, plants, karyotes. Size is a general aspect of cell C
fungi, and animals are eukaryotic cells structure that relates to function. At the lower
(Bolsover et al. 1997). limit, the smallest cells known are bacteria called
Prokaryotic cells are simpler and generally Mycoplasmas, which have diameters between 0.1
smaller than eukaryotic cells and are thought to and 1.0 mm. Eukaryotic cells are typically ten
have evolved first (Fig. 3). Fossils show that pro- times bigger than bacteria.
karyotic organisms antedate by at least 2 billion There are two distinct types (Domains) of pro-
years the first eukaryotes, which appeared some karyotes, the ▶ Bacteria and ▶ Archaea, which
1.5 billion years ago. It is likely that eukaryotes are no more genetically related to each other than
evolved from prokaryotes. The most plausible either group is to the eukaryotes. Both show the
explanation of this process is known as the endo- typical prokaryotic structure where the nucleus
symbiotic theory, first articulated by the Russian and other internal membranes are lacking.
botanist Konstantin Mereschkowski in 1905. The Despite this visual similarity to bacteria, archaea
endosymbiotic theory was advanced and substanti- possess genes and several metabolic pathways
ated with microbiological evidence by Lynn that are more closely related to those of eukary-
Margulis in a 1967 paper, The Origin of Mitosing otes: notably the proteins involved in transcrip-
Eukaryotic Cells. The basis of this theory concerns tion and translation. Other aspects of archaean
the origins of mitochondria and plastids (e.g., chlo- biochemistry are unique, such as their reliance
roplasts), which are organelles of eukaryotic cells. on ether lipids in their cell membranes and cell
According to this theory, these organelles origi- wall. The cell wall of bacteria is always made of
nated as separate prokaryotic organisms that were peptidoglycan, a molecule unique to this group or
taken inside the cell as endosymbionts. Mitochon- organisms. Archaea often have cell walls, but
dria developed from proteobacteria (in particular, peptidoglycan is never present. Thus, the only
Rickettsiales or close relatives) and chloroplasts well defined cellular structures presented by pro-
from cyanobacteria (Margulis 1967). karyotes, the cell membrane and cell wall, are
In addition, prokaryotic cells also lack most chemically quite different in these two groups
other membrane-bound organelles typical of or organisms. Initially, archaea were seen as
eukaryotic cells. In some prokaryotic cells the extremophiles that lived in harsh environments,
plasma membrane is folded inward to form a such as hot springs and salt lakes, but they have
complex of internal membranes (the mesosome) since been found in a broad range of habitats,
along which the relations of cellular respiration such as soils, oceans, and marshlands. Archaea
are thought to take place. Photosynthetic pro- are particularly numerous in the oceans, and the
karyotes contain chlorophyll associated with flat archaea in plankton may be one of the most
membranes called lamellae. On the contrary, abundant groups of organisms on the planet
eukaryotic cells have many types of membrane- (Clark 2005).
bound organelles that partition the cytoplasm into In addition to the plasma membrane, a eukary-
compartments. otic cell has extensive and elaborately arranged
Typically, each prokaryotic cell has a single internal membranes, which partition the cells into
chromosome carrying a full set of genes provid- compartments. The nucleus is surrounded by a
ing it with the genetic information necessary to double membrane, the nuclear membrane, which
operate as a living organism. Each chromosome separates the nucleus from the cytoplasm, but
has 3,000–4,000 genes although some have as allows some communication with the cytoplasm
404 Cell
Capsule
Cell wall
Plasma membrane
Cytoplasm
Ribosomes
Plasmid
Pili
Bacterial Flagellum
Nucleoid (circular DNA)
Nucleus
Nuclear pore
Nuclear envelope Golgi vesicles
Chromatin (golgi apparatus) Lysosome
Nucleolus Centrioles
Ribosomes
Plasma membrane Cytoplasm

Mitochondrion
Peroxisome
Cytoskeleton
Free Ribosomes
Secretory vesicle
Smooth endoplasmic reticulum

Rough endoplasmic reticulum
Flagellum
Cell, Fig. 3 Squematic representation of the cell compo- which carries the archived instructions for operating the
sition. (a) Prokaryotic cells, (b) eukaryotic animal cells. cell. Physiologically they are very similar in many ways.
Despite their apparent differences, both cell types have a For example, the DNA in the two cell types is precisely the
lot in common. They perform most of the same kinds of same kind of DNA, and the genetic code for a prokaryotic
functions, and in the same ways. All are enclosed by cell is exactly the same genetic code used in eukaryotic
plasma membranes, filled with cytoplasm, and loaded cells
with small structures called ribosomes. They have DNA
Cell 405
via nuclear pores. The nucleus contains most of ▶ chloroplasts are the main energy transducers of
the genes in the eukaryotic cell (although some cells. Mitochondria are generally rod-shaped
genes are located in the mitochondria and chlo- organelles bounded by a double membrane. They
roplasts). The genome of eukaryotes usually con- resemble bacteria in their size and shape; it is
sists of 10,000–50,000 genes carried on several thought that mitochondria are indeed evolved
▶ chromosomes. In a cell that is not dividing, from bacteria that took up residence in the prime-
DNA is organized along with proteins forming val ancestor of eukaryotic cells. Like bacteria, C
an irregular network of strands termed chromatin. mitochondria contain a circular molecule of
When the cells began the process of nuclear divi- DNA similar to a bacterial chromosome, although
sion (mitosis), the chromatin coils and condenses much smaller. Mitochondria are the site of most of
into discrete chromosomes containing several the chemical reactions that convert the chemical
thousand genes arranged in a specific linear energy present in inorganic compounds to another
order. Most eukaryotes are diploid, with two cop- form of energy, ATP, that cells can use for work.
ies of each chromosome. Consequently, they pos- These organelles are the sites for cellular respira-
sess at least two copies of each gene. tion, the catabolic process that generates ATP by
Besides nucleus, eukaryotic cells contain a extracting energy from sugars, fats, and other
variety of ▶ organelles, which are subcellular compounds with the help of oxygen. This con-
structures that carry out specific tasks. Some of trasts with bacteria, where the respiration chain is
them are separated from the rest of the cytoplasm located in the cytoplasmic membrane, as no mito-
by membranes but others are not. Many of the chondria are present (Davis et al. 1990).
different membranes of the eukaryotic cell are ▶ Chloroplasts are also membrane-bound
forming an extensive complex of branching organelles that produce and store food materials
tubules, named endoplasmic reticulum (ER) that in algae and plant cells. Chloroplasts contain the
is continuous with the nuclear envelope and per- light-absorbing pigment chlorophyll, along with
meates the cytoplasm. The ER manufactures enzymes and other molecules that function in the
membranes and performs many other biosynthetic photosynthetic production of reduced carbon
functions such as synthesis of lipids, metabolism molecules by trapping light energy. Like mito-
of carbohydrates, and detoxifications of drugs and chondria, chloroplasts contain a circular DNA
toxins. The ER also functions as a system for molecule and are thought to have evolved from
transporting materials from one part of the cell to a photosynthetic bacteria. Chloroplasts also con-
another and perhaps to the outside environment as tain a variety of yellow and orange pigments
well. The Golgi apparatus is a stack of flattened known as carotenoids. Although a unicellular
membrane sacs and associated vesicles that is alga may have only a single large chloroplast,
involved in the secretion of the proteins cells of complex plants may possess 20–100 of
manufactured along the ER. The proteins are these organelles.
released from the ER in sealed-off little vesicles Besides for the presence of chloroplasts, plant
that fuses with the membranes of the Golgi com- cells differ from animal cells in several other
plex. Within the Golgi complex the proteins are ways. Although all cells are limited by plasma
modified in various ways (i.e., adding carbohy- membranes, plant cells are also surrounded by
drates forming glycoproteins). The Golgi appara- ▶ cell walls of cellulose, which limits any change
tus in plant cells produces polysaccharides used to of position and shape. In addition, most plants
construct the cell wall. have one large or several small compartments
Other organelles related to cell metabolism are called vacuoles, used for storing nutrients and
the lysosomes, membrane-bound structures spe- waste products, and certain organelles such as
cialized for digestion that contain hydrolytic centrioles and lysosomes are absent.
enzymes that cells use to digest macromolecules. There are still other structures that support the
About 40 different enzymes have been identified cells, connect them with other cells, and help
in lysosomes. In addition, mitochondria and them move. Cytoskeletal elements, such as
406 Cell
microtubules or microfilaments, give cells their eukaryotes during a time when the oxygen evo-
shape and allow the movement. Centrioles are lution was changing Earth’s environments dra-
tiny organelles that function in nuclear division, matically. Development of chloroplasts may be
usually located within a dense area of cytoplasm, part of the explanation for this temporal
the centrosome. Cells are able to swim or move correlation.
by using their cilia and flagella, which are spe- It seems reasonable to suggest that eukaryotes
cialized arrangements of microtubules (Maton evolved from a single prokaryotic ancestor that
et al. 1997). gradually accumulated greater structural com-
plexity. But evidences show that the eukaryotic
Cell Origin and Evolution cell originated from a symbiotic coalition of mul-
Synthesis and accumulation of biologically rele- tiple prokaryotic ancestors not just one. The the-
vant molecules in the early Earth would have ory of serial endosymbiosis proposes that
been the first step in the path to the primitive mitochondria and chloroplasts were formerly
cells. It is generally assumed that RNA was the small prokaryotes living within larger cells
first information storage molecule and that DNA (Margulis 1967; Davis et al. 1990). The proposed
came later. The primitive cell vaguely resembled a ancestors of mitochondria were aerobic hetero-
bacterium (Bada and Lazcano 2010). trophic prokaryotes, related to alpha-
Life began remarkably early in Earth’s his- proteobacteria, that became endosymbionts. The
tory, and those first organisms were ancestral to proposed ancestors of chloroplasts were photo-
the great diversity of life we observe today. The synthetic prokaryotes related to cyanobacteria.
Earth was formed about 4.5 billion years ago. These ancestors probably gained entry to the
However, the oldest fossils of organisms known host cell as undigested prey or parasites. Evi-
are 3.5 billion years old. These microfossils dences supporting this idea are the structural sim-
resemble certain bacteria that still exist today. ilarity between bacteria, mitochondria, and
For bacteria so complex to have evolved by chloroplasts; both organelles are able to replicate
3.5 billion years ago, it is a reasonable hypothesis by a splitting process reminiscent of binary fis-
that life originated much earlier, when Earth sion in bacteria, and each organelle contains a
began to cool to a temperature at which liquid genome consisting of a single circular DNA mol-
water could exists. The fossil record supports the ecule not associated with histones or other pro-
presumption that prokaryotes were the earliest teins, as in most prokaryotes. Furthermore, the
organisms. Photosynthesis probably evolved organelles contain all the enzymatic equipment
very early also in prokaryotic history. The photo- necessary to transcribe and translate their DNA
synthetic bacteria that generate and release oxy- into proteins. During the evolution from endo-
gen to the atmosphere, named cyanobacteria, symbiont to organelle, the vast majority of the
probably evolved over 2.7 billion years ago. The original bacterial genes has been lost or trans-
accumulation of atmospheric oxygen was gradual ferred to the host nucleus; so the organellar
and had an enormous impact on life since oxygen genomes are very reduced in comparison to
attacks chemical bonds and was toxic for many of their closest free-living counterparts.
the existing prokaryotic groups. Although some
prokaryotic species survived in habitats that
remained anaerobic, a variety of different adap- Basic Methodology
tations to the oxydizing atmosphere evolved,
including cellular respiration, using oxygen to Living cells are composed of many progressively
obtain energy from organic and inorganic smaller components. Most levels of biological
molecules. organization are imperceptible to the human
The oldest fossils of eukaryotes are 2.1–2.7 senses, so that to study them we must make use
billion years old, and look like simple single- of a variety of instruments and indirect tech-
celled algae. This range of time places the earliest niques. Microscopy is the most useful technique
Cell 407
Cell, Fig. 4 (continued)

408 Cell
to study cell structure and related issues. The light allow a magnified view of the sample. The
microscope, gradually improved since Hooke’s resulting image can be detected directly by the
time, uses visible light as the source of illumina- eye. To be viewed with the light microscope,
tion. During the last decades, the development of specimens must be very thin. Single cell organ-
the electron microscope has enabled researchers isms can be observed in vivo. There are important
to study the fine detail, called ultrastructure of limitations to the standard optical microscopy: the
cells. Nowadays, the scanning tunneling electron technique can only image dark or strongly
microscope allows us to study the relationship refracting objects efficiently, diffraction limits res-
between molecules. olution to approximately 0.2 mm, and out of focus
Magnification is the ratio of the size of the light from points outside the focal plane reduces
image to the size of the specimens. Whereas the image clarity. Live cells generally lack sufficient
ordinary light microscope can magnify a struc- contrast to be studied successfully. Internal struc-
ture about 1,000 times, the electron microscope tures of the cell are colorless and transparent.
can magnify up to 250,000 times or even more. The electron microscope uses a beam of elec-
Besides, the electron microscope has far superior trons as a source of illumination instead of light.
resolving power, which is the ability to reveal fine The microscope has a greater resolving power than
detail, and is expressed as the minimum distance a light-powered optical microscope, because it uses
between two points that can be distinguished as electrons that have wavelengths about 100,000
separate and distinct points. times shorter than visible light (photons), and can
Optical and electron microscopy involve the achieve magnifications of up to 1,000,000 times.
diffraction, reflection, or refraction of electromag- The electron microscope uses electrostatic and
netic radiation/electron beams interacting with the electromagnetic “lenses” to control the electron
subject of study, and the subsequent collection of beam and focus it to form an image. Two types of
this scattered radiation in order to build up an electron microscopes in common use are the trans-
image. This process may be carried out by wide- mission electron microscope and the scanning elec-
field irradiation of the sample (e.g., standard light tron microscope (Murphy 2002).
microscopy and transmission electron micros-
copy) or by scanning of a fine beam over the
sample (e.g., confocal laser scanning microscopy See Also
and scanning electron microscopy) (Fig. 4).
Optical or light microscopy involves passing ▶ Archaea
visible light transmitted through or reflected from ▶ Bacteria
the sample through a single or multiple lenses to ▶ Cell Membrane
Cell, Fig. 4 Microscopes and their images. (a) observe the ultrastructure of the nuclei, chloroplasts, and
A confocal microscope creates sharp images of a speci- pyrenoid. (e) Scanning electron microscope (SEM), while
men that would otherwise appear blurred when viewed TEM allows us to study the inner structure of objects
with a conventional optical microscope. This is achieved (tissues, cells, viruses) and SEM is used to visualize the
by excluding most of the light from the specimen that is surface of tissues, macromolecular aggregates, and mate-
not from the microscope’s focal plane. (b) A glia neuron rials. (f) An SEM image of a diatom, brown algae. We can
cell viewed using confocal microscopy. (c) Transmission see the silica theca that surround the cells with their
electron microscope (TEM) uses a high energy electron characteristical patterns specific for each species. (g) The
beam transmitted through a very thin sample to image and scanning tunneling microscope (STM) provides a picture
analyze the microstructure of materials with atomic scale of the atomic arrangement of a surface by sensing corru-
resolution. The electrons are focused with electromag- gations in the electron density of the surface that arise
netic lenses and the image is observed on a fluorescent from the positions of surface atoms. (h) An STM image of
screen, or recorded on film or digital camera. (d) A TEM palladium crystals
image of the green algae Chlamydomonas. We can
Cell Membrane 409
▶ Cellular Theory, History of Definition

▶ Cytoplasm
▶ Eukarya The ▶ cell membrane is the boundary that
▶ Evolution, Biological envelops all cells and provides a semipermeable
▶ Prokaryote barrier for their separation from the extracellular
environment. It is constituted by a 5–8-nm-thick
lipid bilayer – mainly composed of amphiphilic C
phospholipids – in which membrane proteins are
Alberts B, Bray D, Lewis J, Raff M, Roberts K, Watson JD interspersed. The cell membrane is involved in
(1994) Molecular biology of the cell, 3rd edn. Garland different cellular processes including active and
Publ, New York, p 1294 passive traffic of substances, signal transduction,
Bada JL, Lazcano A (2010) The origin of life. In: Ruse M, and cell adhesion and fusion. In addition to the
Travis J (eds) The Harvard companion of evolution.
Harvard University Press, Belknap, Cambridge cell membrane, inside the eukaryotic cytoplasm,
Bolsover SR, Hyams JS, Jones S, Shephard EA, White HA there are membrane-enclosed ▶ organelles.
(1997) From genes to cells. Wiley, New York, p 424 These specialized compartments include the
Campbell NA, Reece JB (2002) Biology, 6th edn. nucleus, mitochondria, plastids, Golgi apparatus,
Benjaming Cummings, San Francisco, p 1247
Clark DP (2005) Molecular biology – understanding the vacuoles, vesicles, lysosomes, peroxisomes, and
genetic revolution. Elsevier Academic Press, Burling- endoplasmic reticulum. Mitochondria, present in
ton, p 783 almost all eukaryotes, and plastids, present in
Davis PW, Solomon EP, Berg LR (1990) Cell structure photosynthetic eukaryotes, are double-membrane
and function. In: The world of biology. Saunders Col-
lege, Fort Worth, pp 79–107 organelles, a feature indicative of their endosym-
Margulis L (1967) On the origin of mitosing cells. J Theor biotic origin. ▶ Bacteria and archaea lack
Biol 14(3):255–274 membrane-enclosed compartments, with certain
Maton A, Hopkins JJ, LaHart S, Warner SQ, Wright M, exceptions such as the magnetosomes present
Jill D (1997) Cells building blocks of life. Prentice
Hall, New Jersey, p 267 within the nucleocytoplasm of magnetotactic
Murphy DB (2002) Fundamentals of light microscopy and bacteria and the cell compartments observed in
electronic imaging. Springer, Heidelberg, p 262 bacteria from the phylum Planctomycetes. Some
▶ virus families – the so-called lipid coated
viruses – are also surrounded by a membrane
during their extracellular phase, derived from
Cell Communication the host cell previously infected.
▶ Quorum Sensing
See Also
Cell Membrane ▶ Amphiphile

▶ Archaea
Carlos Briones ▶ Bacteria
Centro de Astrobiologı́a (CSIC/INTA), Consejo ▶ Cell
Superior de Investigaciones Cientı́ficas, Madrid, ▶ Cell Wall
Spain ▶ Eukarya
▶ Lipid Bilayer
▶ Membrane
Synonyms ▶ Membrane Potential
▶ Organelle
Cytoplasmic membrane; Plasma membrane ▶ Virus
410 Cell Models
detergent was removed, small vesicles formed

Cell Models that contained the original lipids and functional
proteins. Using this method, Racker and his col-
David Deamer leagues were able to reconstitute electron trans-
Department of Chemistry, University of port reactions of mitochondrial and chloroplast
California, Santa Cruz, Santa Cruz, CA, USA membranes.
Similar techniques were soon applied to other
biological structures and functions. For instance,
Keywords Oesterhelt and Stoeckenius (1971) reconstituted
the proton pump of purple membranes isolated
Compartments; Encapsulation; Lipid vesicles; from a species of Halobacterium ( now recog-
Permeability; Replication; Translation nized as archaeon, rather than a bacterium) that
uses the energy of a proton gradient to synthesize
ATP. Racker and Stoeckenius then collaborated
Synonyms to produce a system of reconstituted membrane
vesicles containing both the proton pump of
Artificial cells; Protocell; Synthetic cells Halobacterium halobium and the ATP synthase
of mitochondria (Racker and Stoeckenius 1974).
The hybrid structures could synthesize ATP
Definition using light as an energy source, which strongly
confirmed Peter Mitchell’s chemiosmotic
Cell models are laboratory versions of simple hypothesis that proton gradients could drive
cellular structures that exhibit some of the prop- ATP synthesis (Mitchell 1976).
erties of the living state. They consist of a com- When it was realized that lipid vesicles could
partment, usually microscopic lipid vesicles, with incorporate enzymatic functions, the next step
encapsulated functional polymers such as toward model cells became feasible, in which a
enzymes and nucleic acids. polymerase encapsulated in liposomes would be
able to synthesize a nucleic acid. This was first
attempted by Chakrabarti et al. (1994) and Walde
History et al. (1994) both reporting that encapsulated poly-
nucleotide phosphorylase could synthesize an RNA
The idea that it might be possible to assemble homopolymer from its substrate, in this case ADP.
model cells can be traced back to 1965, when
Alec Bangham discovered that phospholipids
spontaneously self-assemble into closed compart- Overview
ments (now called liposomes) when dispersed in
aqueous phases (see review by Bangham 1993). The point of this brief history is that relatively
The boundary of such compartments is a ▶ lipid complex biological functions can be
bilayer that is relatively permeable to water and reconstituted by ▶ self-assembly of their dis-
small molecules like water but much less perme- persed components, so it is reasonable to consider
able to ionic solutes such as sodium and potas- the possibility that similar techniques might
sium ions. Efraim Racker took the next step allow artificial cells to be fabricated under labo-
toward cell models when he and his coworkers ratory conditions. If this turns out to be possible,
showed that it was possible to use detergents such perhaps it will help define “life” and even eluci-
as deoxycholic acid to disperse membranous com- date the major steps that led to the origin of
ponents of cells (see Racker 1970). When the cellular life nearly four billion years ago.
Cell Models 411
Basic Methodology Key Research Findings
What would such a system do? This Several research groups are beginning to study
question can be answered by listing the prop- systems of genetic and catalytic molecules that
erties and functions of model cells that could are steps toward fulfilling this list of properties.
conceivably be assembled in the laboratory. For instance, Mansy et al. (2008) demonstrated
For the purposes of this list, it is assumed that it was possible to encapsulate a short DNA C
that all the nutrients and energy needed for template in fatty acid vesicles, then add activated
growth and replication will be provided so nucleotides outside. The nucleotides were suffi-
that a complex metabolism will not be ciently permeable to enter the vesicle interior and
required: support the sequence-dependent elongation of the
DNA. In a related advance, Lincoln and Joyce
1. Self-assembly of lipid molecules generates (2009) developed a pair of ribozymes, each of
cellular compartments defined by boundary which could catalyze the synthesis of the other by
membranes. a ligation reaction that joined two smaller
2. Macromolecules are encapsulated in the non-catalytic oligonucleotides. This system is
compartments, yet smaller substrate mole- not encapsulated and does not replicate a com-
cules can cross the membrane barrier. plete base sequence but illustrates in principle
3. The macromolecules have the potential to how an evolving system of paired ribozymes
grow by polymerizing the substrate could function. The Rasmussen Group in Den-
molecules. mark has stepped away from biologically
4. The membrane itself can grow by addition of inspired molecular systems and is attempting to
lipid molecules. assemble a very different version of model cells
5. Some of the encapsulated polymers are cata- (DeClue et al. 2009). In their system, the reac-
lysts that can speed the growth process, and tions occur on the surface of vesicles, rather than
the catalysts are reproduced during growth the interior, thereby bypassing the requirement
by polymerization. for membrane transport of the nutrients. Further-
6. Genetic information is contained in the more, they are attempting to drive the polymeri-
sequence of monomers in a second set of zation reaction by an input of light energy, rather
polymers and is used to direct the growth of than supplying activated monomers.
catalytic polymers. Another approach to model cells is to encap-
7. The catalytic polymers catalyze the polymer- sulate ribosomes and translation systems in lipid
ization of the genetic molecules. vesicles. Luigi Luisi and his coworkers at the
8. Following a certain amount of growth, the Eidgenössische Technische Hochschule in
membrane-bounded system of macromole- Zurich, Switzerland, made the first attempt to
cules divides into smaller structures, each assemble a translation system in lipid vesicles
containing the catalysts and copies of genetic by encapsulating ribosomes and an RNA homo-
information. polymer that codes for phenylalanine
9. Genetic information is passed between (Oberholzer et al. 1995). The phenylalanine was
generations by duplicating the gene attached to transfer RNA so that it was ready to be
sequences and sharing them between used by ribosomes for peptide synthesis. How-
daughter cells. ever, the lipid bilayer was impermeable to the
10. Occasional mistakes (mutations) occur dur- transfer RNA, so peptide bond formation was
ing replication or transmission of genetic limited to the small number of tRNA-amino
information so that the system can evolve acid complexes that were encapsulated within
through selection. the vesicles. This limitation makes the point that
412 Cell Models
model cells need to have some way to transport bilayer; otherwise, the vesicles have no access
nutrients inward across their boundary to external sources of nutrients and energy.
membrane. A whole set of regulatory processes must be in
Noireaux and Libchaber (2004) reported an place so that all of these functions are coordi-
elegant solution to the permeability problem. nated. Finally, when the vesicles grow to approx-
They disrupted E. coli cells and captured samples imately twice their original size, there must be a
of the bacterial cytoplasm in lipid vesicles. The way for them to divide into daughter cells that
samples included ribosomes, transfer RNAs, and share the original genetic information.
the hundred or so other components required for It seems impossible that the first forms of life
protein synthesis. The researchers then chose two sprang into existence with such a complex system
genes to translate, one for green fluorescent proof interacting molecules. There must have been
tein (GFP), a marker for protein synthesis, and a something simpler, a kind of scaffold life that was
second gene for a pore-forming protein called left behind in the evolutionary process leading to
alpha hemolysin. If the system had worked as today’s life. Can we reproduce that scaffold? This
planned, the GFP would have accumulated in is the challenge for research on model cells and
the vesicles as a visual marker for protein synthe- the origin of life.
sis and the hemolysin would have allowed exter-
nally added “nutrients” in the form of amino
acids and ATP to cross the membrane barrier Applications
and supply the translation process with energy
and monomers. The system was functional, and A functioning system of artificial cells will rep-
the newly synthesized hemolysin allowed synthe- resent a major breakthrough in biotechnology. At
sis of GFP to continue for as long as four days. present, the pharmaceutical industry must use
The GFP in the vesicles was monitored by its recombinant DNA techniques and bacterial cul-
green fluorescence. A more recent example of a tures to synthesize protein products. Model cells
model cell system is the encapsulated “genetic are simplified versions of bacterial cells, and if
cascade” fabricated by Ichihashi et al. (2010), in they can be designed to produce a desired protein
which a gene on a plasmid was transcribed by in large quantities, the result would be a much
RNA polymerase to mRNA, which in turn more efficient, flexible, and inexpensive system
directed the synthesis of green fluorescent for producing important therapeutic agents. See
protein. review by Pohorille and Deamer (2002).
The model cells containing ribosomes clearly
demonstrate one fundamental property of life:
they can use genetic information to synthesize a Future Directions
protein. Although they can grow by synthesizing
one or more specific proteins, no other cellular One promising approach to model cells is
components are produced. To approach the defi- suggested by the results reported by Lincoln and
nition of a living system, the vesicles would need Joyce (2009). It is possible that a pair of
to incorporate genetic information required for a ribozymes will be found that can catalyze their
hundred or more different proteins and RNA spe- own complete synthesis using genetic informa-
cies, over half of which are the components of the tion encoded in their base sequences. If the
ribosomes themselves. They would need genes ribozymes could then function in a membrane-
for polymerase enzymes so that the DNA could bounded compartment using nucleotides present
be replicated as part of the growth process, and a in the external medium, the system could rightly
way for lipid to be synthesized, because the mem- be claimed to have the essential properties that
branous boundary must grow to accommodate are lacking so far in artificial cell models: repro-
the internal growth. Transport proteins must be duction of the catalysts and genetic information
synthesized and incorporated into the lipid in a cellular compartment.
Cell Wall 413
See Also Oberholzer TR, Wick R, Luisi PL, Biebricker CK

(1995) Protein expression in liposomes. Biochem
Biophys Res Commun 207:250
▶ Cell Oesterhelt D, Stoeckenius W (1971) Rhodopsin-like pro-
▶ Lipid Bilayer tein from the purple membrane of Halobacterium
▶ Nucleic Acids halobium. Nat New Biol 233:149–152
▶ Protocell Pohorille A, Deamer DW (2002) Artificial cells: prospects
for biotechnology. Trends Biotechnol 20:123
▶ Self-Assembly Racker E (1970) Membranes of mitochondria and chloro-
C
plasts. Van Nostrand Reinhold, New York
Racker E, Stoeckenius W (1974) Reconstitution of purple
membrane vesicles catalyzing light-driven proton
uptake and adenosine triphosphate formation. J Biol
References and Further Reading Chem 249:662–663
Schrum JP, Ricardo A, Krishnamurthy M, Blain JC,
Bangham AD (1993) Liposomes: the Babraham connec- Szostak JW (2009) Efficient and rapid template-
tion. Chem Phys Lipids 64:275–285 directed nucleic 258 acid copying using 20 -amino-20 ,3-
0
Chakrabarti A, Breaker RR, Joyce GF, Deamer DW -dideoxyribonucleoside-50 -phosphorimidazolide
(1994) Production of RNA by a polymerase protein monomers. J Am Chem Soc 131:14560–14570
encapsulated within phospholipid vesicles. J Mol Sunami T, Kita H, Hosoda K, Matsuura T, Suzuki H,
Evol 39:555–559 Yomo T (2009) Detection and analysis of protein
Deamer D, Szostak J (eds) (2010) Origins of life. Cold synthesis and RNA replication in giant liposomes.
Spring Harbor Press, Woodbury Methods Enzymol 464:19–30
Deamer DW, Dworkin JP, Sandford SA, Bernstein MP, Szostak JW, Bartel DP, Luisi PL (2001) Synthesizing life.
Allamandola LJ (2002) The first cell membranes. Nature 409:387–390
Astrobiology 2:371–382 Walde P, Goto A, Monnard P-A, Wessicken M, Luisi PL
DeClue MS, Monnard PA, Bailey JA, Maurer SE, Collis (1994) Oparin’s reactions revisited: enzymatic synthe-
GE, Ziock HJ, Rasmussen S, Boncella JM sis of poly(adenylic acid) in micelles and self-
(2009) Nucleobase mediated, photocatalytic vesicle reproducing vesicles. J Am Chem Soc 116:7541–7547
formation from an ester precursor. J Am Chem Soc Zhu TF, Szostak JW (2009) Coupled growth and division
131:931–933 of model protocell membranes. J Am Chem Soc
Hanczyc MM, Szostak JW (2004) Replicating vesicles as 131:5705–5713
models of primitive cell growth and division. Curr
Opin Chem Biol 8:660–664
Ichihashi N, Matsuura T, Kita H, Sunami T,
Suzuki H, Yomo T (2010) Constructing partial
models of cells. In: Deamer D, Szostak J (eds) Origins
of life. Cold Spring Harbor Laboratory Press,
Cell Motility
Woodbury, NY
Lincoln TA, Joyce GF (2009) Self-sustained replication of ▶ Motility
an RNA enzyme. Science 323:1229–1232
Luisi PL (2006) The emergence of life: from chemical
origins to synthetic biology. Cambridge University
Press, Cambridge
Mansy SS, Schrum JP, Krishnamurthy M, Tobé S, Treco Cell Wall
DA, Szostak JW (2008) Template-directed synthesis
of a genetic polymer in a model protocell. Nature
Felipe Gomez
454:122–125
Mitchell P (1976) Possible molecular mechanisms of the Centro de Astrobiologı́a (CSIC/INTA), Instituto
protonmotive function of cytochrome systems. J Theor Nacional de Técnica Aeroespacial, Torrejón de
Biol 62:327–367 Ardoz, Madrid, Spain
Monnard PA, Luptak A, Deamer DW (2007) Models of
primitive cellular life: polymerases and templates in
liposomes. Philos Trans R Soc Lond B Biol Sci
Morowitz HJ (2009) Beginnings of cellular life. Yale
University Press, New Haven
Noireaux V, Libchaber A (2004) A vesicle bioreactor as a
The cell wall is an external layer that surrounds
step toward an artificial cell assembly. Proc Natl Acad some types of ▶ cells. It has structural, protective,
Sci U S A 101:17669–17674 and functional roles (filtering capacities for the
414 Cell, Minimal
selective uptake of substances for cell metabolism). Overview

It is located outside the cell membrane. The cell
wall is found in bacteria, plants, fungi, algae, and A minimal living cell must still retain all essential
some Archaea. The main functions of cell walls are genes needed to perform all essential functions, i.e.,
(1) to provide tensile strength and limited plasticity a minimal genome. However, there is no concep-
to the cell, (2) to provide mechanical support, tual or experimental support for the existence of
(3) cutinized, it is used to prevent water loss, one form of minimal cell. Different essential func-
(4) to provide mechanical protection, and (5) to tions can be defined depending on the environment,
contribute to cell-cell communication. The struc- and numerous versions of minimal genomes can be
ture of the cell wall differentiates Gram-positive envisaged to perform such functions even in the
from ▶ Gram-negative bacteria, a property of same conditions (Koonin 2003). Computational
important phylogenetic value. and experimental studies (mostly bacterium cen-
tered) to define the characteristics of a minimal cell
(reviewed in Feher et al. 2007; Moya et al. 2009)
converge in the conclusion that different minimal
See Also
hypothetically viable cells can be conceived,
possessing a universal genetic machinery
▶ Cell
supported by a diversity of minimal ecologically
▶ Cell Membrane
dependent metabolic charts (Gil et al. 2004).
▶ Gram-Negative Bacteria
Defining the minimal genome does not fully
▶ Gram-Positive Bacteria
define a minimal cell. Several levels of complex-
▶ Peptidoglycan
ity (topological genome architecture,
▶ Protoplast
transcriptome, proteome, metabolome,
interactome) that are not taken into account in
minimal genome studies (Fisunov et al. 2011) can
be explored through theoretical computer models
Cell, Minimal based on the massive amount of molecular and
cellular biology data available (Di Ventura
Rosario Gil et al. 2006; Karr et al. 2012; Shuler et al. 2012).
Institut Cavanilles de Biodiversitat i Biologia However, to date, no single computational
Evolutiva, Universitat de València, Paterna method can fully explain complex phenotypes
(València), Spain in terms of molecular components and their inter-
actions, and it is difficult to determine and model
all parameters involved (Karr et al. 2012).
Keywords Nowadays, one of the main goals of synthetic
biology is the generation of minimal cells, to
Bottom-up approach; Minimal genome; Syn- improve our basic understanding of living sys-
thetic biology; Top-down approach tems and to be used as containers for introducing
genetic modules to customize cells for biotech-
nological purposes (Moya et al. 2009). There are
Definition two complementary and alternative ways to do
so, namely, top-down and bottom-up approaches.
A minimal ▶ cell is a hypothetical biological The top-down strategies, also called ▶ genome-
system that possesses only the necessary and driving cell engineering (O’Malley et al. 2008),
sufficient attributes to be considered alive. There- starts from simple modern cells and intends to
fore, it must be able to maintain its own structures simplify them further through the removal of
(homeostasis), self-reproduce, and evolve in a nonessential genetic elements. The bottom-up
supportive, protected, and stable environment. approaches aim at constructing the simplest
Cellular Automata 415
system that could be considered alive from Karr JR, Sanghvi JC, Macklin DN, Gutschow MV, Jacobs
scratch (Luisi 2002), by putting together the JM, Bolival B Jr, Assad-Garcia N, Glass JI, Covert
MW (2012) A whole-cell computational model pre-
essential nonliving components, a self- dicts phenotype from genotype. Cell 150:389–401
replicating nucleic acid, a metabolic machinery, Koonin EV (2003) Comparative genomics, minimal gene-
and an encapsulating structure (Stano and Luisi sets and the last universal common ancestor. Nat Rev
2011), although no such system has been Microbiol 1:127–136
Luisi PL (2002) Toward the engineering of minimal living
built yet. cells. Anat Rec 268:208–214
C
Research based on extant cells to define a Moya A, Gil R, Latorre A, Pereto J, Pilar Garcillan-
hypothetical minimal cell is not related with the Barcia M, de la Cruz F (2009) Toward minimal bacte-
study of the origin of life, because it is impossible rial cells: evolution vs. design. FEMS Microbiol Rev
33:225–235
to identify which one of the putative alternatives O’Malley MA, Powell A, Davies JF, Calvert J (2008)
was adopted by primitive cells. In contrast, some Knowledge-making distinctions in synthetic biology.
bottom-up research programs are motivated by Bioessays 30:57–65
the possibility of better understanding this topic, Pereto J, Catala J (2007) The renaissance of synthetic
biology. Biol Theor 2:128–130
by recreating some of the steps that pre-cellular Shuler ML, Foley P, Atlas J (2012) Modeling a minimal
systems might have followed under primitive cell. Methods Mol Biol 881:573–610
Earth conditions (Pereto and Catala 2007). Stano P, Luisi PL (2011) On the construction of minimal
cell models in synthetic biology and origins of life
studies. In: Koeppl H, Setti G, di Bernardo M,
Densmore D (eds) Design and analysis of biomolecu-
See Also lar circuits. Springer, Dordrecht, pp 337–368
▶ Cell
▶ Genome
▶ Genome, Minimal Cellular Automata
Marco Tomassini
References and Further Reading Information Systems Department, University of
Lausanne, Lausanne, Switzerland
Bersini H (2010) Minimal cell: the computer scientist
point of view. In: Gargaud M, Lopez-Garcia P, Martin
H (eds) Origin and evolution of life: an astrobiology
perspective. Cambridge University Press, Cambridge, Keywords
Chap 4
Brochier C (2010) Minimal cell: the biologist point of Automata; Complex systems; Discrete dynamics;
view. In: Gargaud M, Lopez-Garcia P, Martin
H (eds) Origin and evolution of life: an astrobiology
Simulation
perspective. Cambridge University Press, Cambridge,
Chap. 3
Di Ventura B, Lemerle C, Michalodimitrakis K, Serrano Synonyms
L (2006) From in vivo to in silico biology and back.
Nature 443:527–533
Feher T, Papp B, Pal C, Posfai G (2007) Systematic Tessellation automata
genome reductions: theoretical and experimental
approaches. Chem Rev 107:3498–3513
Fisunov GY, Alexeev DG, Bazaleev NA, Ladygina VG,
Galyamina MA, Kondratov IG, Zhukova NA,
Definition
Serebryanova MV, Demina IA, Govorum VM
(2011) Core proteome of the minimal cell: compara- Cellular automata (CA) are dynamical systems in
tive proteomics of three mollicute species. PLoS One which space and time are discrete. A cellular
6:e21964
Gil R, Silva FJ, Pereto J, Moya A (2004) Determination of
automaton consists of an array of cells, each of
the core of a minimal bacterial gene set. Microbiol Mol which can be in one of a finite number of possible
Biol Rev 68:518–537 states, updated synchronously in discrete time
416 Cellular Automata
Cellular Automata, Table 1 Parity rule table. CNESW denotes the current states of the center, north, east, south, and
west cells, respectively. S next is the state of the central cell state at the next time step
CNESW Snext CNESW Snext CNESW Snext CNESW Snext
00000 0 01000 1 10000 1 11000 0
00001 1 01001 0 10001 0 11001 1
00010 1 01010 0 10010 0 11010 1
00011 0 01011 1 10011 1 11011 0
00100 1 01100 0 10100 0 11100 1
00101 0 01101 1 10101 1 11101 0
00110 0 01110 1 10110 1 11110 0
00111 1 01111 0 10111 0 11111 1
steps, according to a local, identical interaction the states of its four neighbors in the N, E, S, and
rule. The state of a ▶ cell at the next time step is W directions is odd and is assigned a state of 0 if
determined by its own current state and the cur- the parity is even. There are 32 different combi-
rent states of a surrounding neighborhood of cells nations for the states of the neighbors, including
(Wolfram 1994). the central cell itself. The rule table consists of
entries of the form.
Overview
0
Cellular automata were originally conceived by
Ulam and von Neumann in the 1940s to provide a 1 1 0 1
formal framework for investigating the behavior
of complex, extended systems (von Neumann
1966). In particular, von Neumann asked whether 1
we could use purely mathematical-logical con-
siderations to discover the specific features of This means that if the current state of the cell is
automata that make them formally analogous 1 and the states of the north, east, south, and west
with self-constructing and self-replicating bio- cells are 0, 0, 1, 1, respectively, then the state of
logical systems. the central cell at the next time step will be
Thanks to their simplicity and appeal, over the 1 (because three bits in the neighborhood are in
years, CA have been applied to the study of gen- state 1). The rule is completely specified by the
eral phenomenological aspects of the world, rule table given in Table 1. Figure 1 demonstrates
including communication, computation, construc- patterns that are produced by the parity CA.
tion, growth, reproduction, competition, and evo- The simplest CA are one dimensional with only
lution. CA have also been used successfully as an two possible states per cell and a neighborhood
easy way to program models for studying phenom- constituted of the cell itself and its immediate
ena of interest in several scientific fields, including right and left neighboring cells. Figure 2 shows an
physics, biology, and computer science. example of the time evolution of an elementary CA.
More formally, a cellular automaton A is a
quadruplet:
Basic Methodology
As an example, let us consider the parity rule for A ¼ ðS, G, d, f Þ

a two-state, five-neighbor, two-dimensional CA.
Each cell is assigned a state of 1 at the next time where S is a finite set of states, G is the cellular
step if the combined parity of its current state and neighborhood, d Z + is the dimension of A, and
Cellular Automata 417
Cellular Automata, Fig. 1 Patterns produced by the represent cells in state 1. a After 30 time steps (t = 30),
parity rule, starting from a 20 20 rectangular pattern. b t = 60, c t = 90, d t = 120
White squares represent cells in state 0, and black squares
f is the local cellular interaction rule, also referred In this case, the domain of f is the set of all 23
to as the transition function. triplets, which gives rise to 28 = 256 distinct
Given the position of a cell i, i Z d, in a elementary rules (Wolfram 1994). For finite-
regular d-dimensional uniform lattice, or grid size grids, spatially periodic boundary conditions
(i.e., i is an integer vector in a d-dimensional are frequently assumed, resulting in a circular
space), its neighborhood G is defined by grid. An example of an elementary rule has
been shown in Fig. 2.
Gi ¼ fi, i þ r1 , i þ r2 , . . . , i þ rn g; For a CA of size N, a configuration of the grid
at time t is defined as
where n is a fixed parameter that determines the CðtÞ ¼ ðs0 ðtÞ, s1 ðtÞ, . . . , sN1 ðtÞÞ;
neighborhood size and rj is a fixed vector in the
d-dimensional space. where s i (t) S is the state of cell i at time t. The
The local transition rule f progression of the CA in time is then given by the
iteration of the global mapping F:
f : Sn ! S
F : CðtÞ ! Cðt þ 1Þ, t ¼ 0, 1, . . .
maps the state si S of a given cell i into another
state from the set S, as a function of the states of through the simultaneous application in each cell
the cells in the neighborhood G i. of the local transition rule f. The global dynamics
Consider a one-dimensional CA with only two of the CA can be described as a directed graph,
states S = {0,1}. In this case, f is a function f: referred to as the CA’s phase space (Wolfram
{0,1}n ! {0,1} and the neighborhood size n is 1994).
usually taken to be n = 2r + 1 such that
si ðt þ 1Þ ¼ f ðsir ðtÞ, . . . , si ðtÞ, . . . , siþr ðtÞÞ; Key Research Findings

where r Z + is a parameter, known as the Cellular automata have been proved to be univer-
radius, representing the standard sal computing devices and, as such, they can
one-dimensional cellular neighborhood. Consid- compute any computable function (Wolfram
ering the r = 1 case, one obtains the so-called 1994). The question of whether cellular automata
elementary CA, for which the neighborhood size can model not only general phenomenological
is n = 3: aspects of our world but also directly model the
laws of physics themselves was raised by Fredkin
f : f0, 1g3 ! f0, 1g, si ðt þ 1Þ and Toffoli (1982). A primary theme of this
¼ f ðsi1 ðtÞ, si ðtÞ, siþ1 ðtÞÞ: research is the formulation of computational
418 Cellular Automata
phenomenon at hand may give rise to a global

behavior that approaches that of the original sys-
tem. Examples of this range from ▶ chemotaxis, to
snow transport by the wind, and to car traffic, just
to name a few (Chopard and Droz 1998). Of
course, how to choose the local cellular automata
rules in order to produce the desired global behav-
ior is a hard problem. It can be approached by trial
and error or by heuristics such as evolutionary
algorithms (Sipper 1997; Crutchfield et al. 2003)
in which good rules evolve out of a population of
possible candidate rules by the Darwinian princi-
ples of variation and selection. CA have thus been
used as a simple and easy to implement model for
studying phenomena of interest in several scien-
tific fields, including physics, biology, engineer-
ing, and computer science.
Cellular Automata, Fig. 2 A one-dimensional elemen-
tary CA is shown, where the horizontal axis depicts the
configuration at a certain time t and the vertical axis Future Directions
depicts successive time steps (increasing down the page).
The system starts in a randomly generated configuration of
zeroes and ones In summary, CA suggest a new approach in
which complex behavior arises in a bottom-up
manner from nonlinear, spatially extended, local
models of physics that are information preserving
interactions and provide a simple and useful
and thus retain one of the most fundamental fea-
model for many complex systems arising in the
tures of microscopic physics, namely, reversibil-
sciences and in engineering.
ity (Fredkin and Toffoli 1982; Margolus 1984;
CA exhibit massive parallelism, locality of
Toffoli 1980). This approach has been used to
cellular interactions, and simplicity of basic hard-
provide extremely simple models of common
ware components. As a consequence, in recent
differential equations of physics, such as the
years, there has been a growing interest in the
heat and wave equations (Toffoli 1984) and the
utilization of CA as actual embedded computing
Navier-Stokes equation (Frisch et al. 1986). CA
devices, e.g., in low-level vision, pseudorandom
also provide a useful model for a branch of
number generation, and cryptography
dynamical systems theory which studies the
(Chaudhuri et al. 1997). A trend that could
emergence of well-characterized collective phe-
become important in the future is the search for
nomena, such as order, turbulence, chaos, sym-
practical quantum computing CA devices, a con-
metry breaking, and fractality, in discrete systems
cept that is potentially capable of revolutionizing
(Chopard and Droz 1998; Vichniac 1984).
computation and its applications; quantum cellu-
lar automata belong to this class (Pérez-Delgado
and Cheung 2007).
Applications
For engineers and scientists, cellular automata are See Also

a particularly useful modeling device when the
phenomenon to be studied does not lend itself to ▶ Autopoiesis
a clear mathematical model such as those ▶ Biological Networks
represented by differential equations. In this case, ▶ Chaotic Region
simple local rules that are motivated by the ▶ Self-Replication
Cellular Theory, History of 419
References and Further Reading History
Chaudhuri PP, Chowdhury DR, Nandi S, Chattopadhyay The cellular theory was formulated after more
S (1997) Additive cellular automata: theory and appli-
than one and a half century of microscopic obser-
cations. IEEE Computer Society, Los Alamitos
Chopard B, Droz M (1998) Cellular automata modeling of vations. Indeed, the invention of microscope dur-
physical systems. Cambridge University Press, ing the seventeenth century offered new
Cambridge investigations to naturalists. For example, Robert C
Crutchfield JP, Mitchell M, Das R (2003) Evolutionary
Hooke (1635–1703) in his Micrographia (1865),
design of collective computation in cellular automata.
In: Crutchfield JP, Schuster P (eds) Evolutionary Antoni van Leeuwenhoek (1632–1723), and Jan
dynamics: exploring the interplay of selection, acci- Swammerdam (1737–1780) observed micro-
dent, neutrality, and function. Oxford University Press, scopical objects: individual entities, often
Oxford, pp 361–411
named animalcules, and parts of organisms.
Fredkin E, Toffoli T (1982) Conservative logic. Int
J Theor Phys 21:219–253 However, they did not have concepts to consider
Frisch U, Hasslacher B, Pomeau Y (1986) Lattice-gas all these new observations. For example, it is
automata for the Navier-Stokes equation. Phys Rev important to notice that when Hooke used the
Lett 56:1505–1508
word cell to name some little spaces that he had
Margolus N (1984) Physics-like models of computation.
Phys D 10:81–95 observed in vegetable organism, he did not
Pérez-Delgado C, Cheung D (2007) Local unitary quan- expect to conceptualize anything, and his inten-
tum cellular automata. Phys Rev A 76:032320 tion was only descriptive.
Sipper M (1997) Evolution of parallel cellular machines:
During all the eighteenth century, microscopic
the cellular programming approach. Springer-Verlag,
Heidelberg observations were more and more accurate. They
Toffoli T (1980) Reversible computing. In: De showed a broad diversity of animalcules and
Bakker JW, Van Leeuwen J (eds) Automata, microscopic structures in living beings. At the
languages and programming. Springer-Verlag, Berlin,
beginning of the nineteenth century, some theo-
pp 632–644
Toffoli T (1984) Cellular automata as an alternative to ries claimed that there could be a unity in the
(rather than an approximation of) differential equa- microscopic structure. The French botanist
tions in modeling physics. Phys D 10:117–127 Charles-François Brisseau de Mirbel
Vichniac G (1984) Simulating physics with cellular
(1776–1854) suggested that plants were consti-
automata. Phys D 10:96–115
Von Neumann J (1966) Theory of self-reproducing tuted by a set of membranes with a lot of pores.
automata. University of Illinois Press, Illinois, Edited A few years later, René Joachim Henri Dutrochet
and completed by Burks AW (1776–1847), who had discovered the osmotic
Wolfram S (1994) Cellular automata and complexity.
phenomenon, claimed that cells constituted
Addison-Wesley, Reading
plants. However, in the wall of these cells, there
could be some little globules that could be the
fundamental entities. Then, François-Vincent
Raspail (1794–1878) (one of the inventors of
Cellular Theory, History of the microscopic colorations) considered that liv-
ing beings were made of globules that were to be
Stéphane Tirard formed in the wall of other globules which would
Centre François Viète d’Histoire des Sciences fit into each other. In that period, Lorenz
et des Techniques EA 1161, Faculté des Okenfuss (Oken) (1779–1851) claimed that liv-
Sciences et des Techniques de Nantes, Nantes, ing beings were a synthesis of infusorians. All
France these proposals have in common the fact that they
are conceptions about the possibility of a micro-
scopic and universal structure.
Keywords In 1838, the German botanist Matthias
Schleiden (1804–1881) claimed that vegetable
Chromosomes; Globules organisms were composed of cells in which
420 Cenancestor
there was a systematic structure, the cytoblast Definition

(which will be later named nucleus). Besides,
Schleiden suggested that cells were the result of The cenancestor is the most recent ancestral spe-
the accumulation of a liquid, the cytoblastem, cie from which all extant living species have
between the cytoblast and the membrane. His evolved. The idea that all present-day life is
colleague, Theodor Schwann (1810–1882), a related by common ancestry was already
specialist of animal physiology, generalized this suggested by Charles Darwin in The Origin of
theory to animals and indicated that the Species (1872). Nowadays, this hypothesis is
cytoblastem came from the interstitial fluid. strongly supported by the similarities at the bio-
During the 1850s, Robert Remak (1815–1865) chemical and molecular levels of all organisms
and Rudolph Virchow (1821–1902) indepen- belonging to the three domains of life, i.e.,
dently (Remak in 1855 and Virchow in Archaea, Bacteria, and Eukarya. The nature of
1855–1858) asserted that every cell was the result this ancestral entity continues to be a matter
of the division of a previous cell. From this time of debate. Its level of complexity (in number of
forth, the concept of cell has become central in genes), the chemical nature of its genome
biology. (whether DNA or RNA), or its prokaryotic-like
During the end of the nineteenth century, the nature have been discussed along other aspects of
progress of microscopy led to observations of its biology.
chromosomes and to the description of mitosis
(Fleming 1882) and meiosis (Boveri, Hertwig
1887–1892).
See Also
▶ Common Ancestor
See Also ▶ Darwin’s Conception of the Origins of Life
▶ Domain (Taxonomy)
▶ Animalcules ▶ Homology
▶ Cell ▶ Last Universal Common Ancestor
▶ Protoplasmic Theory of Life ▶ Phylogeny
▶ Spontaneous Generation, History of

Centaurs (Asteroids)
Duchesneau F (1987) Genèse de la théorie cellulaire.
Bellarmin-Vrin, Paris
Therese Encrenaz
Meudon, France
Cenancestor
Definition
Luis Delaye
Departamento de Ingenierı́a Genética, The Centaurs are outer ▶ asteroids whose orbits
CINVESTAV-Irapuato, Irapuato, Gto, Mexico are mostly confined between those of Jupiter and
Neptune. Due to giant planets’ perturbations,
these objects have transient orbits with typical
Synonyms lifetimes of a few million years. There are a few
tens of Centaurs presently known, among them
Last universal common ancestor (2060) Chiron (also named 95P/Chiron),
Ceres 421
(5145) Pholus, and (10199) Chariklo, the biggest

Centaur found to date with a diameter of 260 km. Ceres
Saturn’s satellite Phoebe is believed to be a cap-
tured Centaur. This population appears to be Ralf Jaumann
intermediate between asteroids and ▶ comets, as German Aerospace Center (DLR), Institute of
any Centaur coming close enough to the Sun is Planetary Research, Berlin, Germany
expected to show cometary activity. C
Keywords
See Also
Asteroid; Dwarf planet
▶ Asteroid
▶ Comet
▶ Kuiper Belt Definition
▶ Trans-neptunian Object
Discovered in 1801 by Giuseppe Piazzi, 1 Ceres
is the largest and one of the oldest and most intact
objects in the asteroid belt, cataloged by the IAU
as a ▶ dwarf planet in 2006. Ceres orbits the
Center of Mass ▶ Sun at a distance of 2.76 AU and differs from
any other ▶ asteroid visited so far. Its surface
▶ Barycenter seems to be covered with water ice and ▶ clay,
a hydrated ▶ rock alteration product, and might
have regions covered with water ice frost. This is
consistent with thermal models making Ceres an
icy object that has been subject to differentiation
Center of Mass Velocity and hydrothermal activity and that might host a
liquid subsurface layer even today.
Daniel Rouan
Meudon, France Overview
(1) Ceres (Fig. 1) is an oblate spheroid with an

Synonyms equatorial radius of 487 2 km and a polar
radius of 455 2 km, a mass of (9.43 0.05)
Barycenter velocity 1020 kg, a density of 2,077 kg/m3, a rotation
period of 9 h, and an orbital period of
4.6 years (Thomas et al. 2005; McCord and
Definition Sotin 2005). The physical properties of Ceres
are consistent with a rocky core and a thick
The center of mass velocity of a system of mas- outer ▶ mantle of ▶ water ice and possibly even
sive points or objects is the velocity of the point a global ocean of water beneath that ice (McCord
where the resultant force of gravitational attrac- and Sotin 2005; McCord et al. 2011). Its surface
tion acts. The system’s whole mass can be con- temperatures vary with latitude from 130 K at the
sidered to be concentrated at this point, for the poles to 180 K at the equator, reaching a maxi-
purpose of calculations. The motion of the center mum of 235 K (cf. Li et al. 2006; Castillo-Rogez
of mass of an object in free fall is the same as the and McCord 2009; McCord et al. 2011). This is
motion of a point object located there. greater than any creep temperature for known icy
422 Ceres
ammoniated saponite or montmorillonite (King

et al. 1992; Rivkin et al. 2006), implying temper-
atures since formation of less than 400 K. In sum-
mary all available chemical data suggest Ceres to
be an object that has experienced extensive aque-
ous alteration by CO2-bearing fluids, with a pow-
dery regolith and little compositional variations
across the surface (Rivkin et al. 2011).
Models constrained by the thermal and com-
positional conditions demonstrate that Ceres
almost certainly differentiated, involving pro-
cesses such as the formation of a silicate core, a
liquid water mantle, and a solid ice crust and
crustal evolution by tectonics and probable
▶ cryovolcanism (Castillo-Rogez and McCord
2009; McCord et al. 2011). Ceres’ relatively
thin ▶ hydrosphere might imply a connection
between endogenic activity and features on the
Ceres, Fig. 1 Dawn imaging Ceres during optical naviga- surface, indicating some remarkably recent pro-
tion on April 14 and 15, 2015 from a distance of 22,000km
showing the north polar region. (NASA / JPL-Caltech / cesses and some astrobiological potential.
UCLA / MPS / DLR / IDA) Ceres will be explored in detailed by the Dawn
mission (launched 2007) (Russell et al. 2007;
Russell and Raymond 2011) that will orbit
compositions, implying Ceres’ icy shell to be in Ceres for 9 months in 2015.
▶ hydrostatic equilibrium. Surface ▶ albedo
varies from 0.04 to 0.09 (Li et al. 2006;
Rivkin et al. 2011), which is suggestive of surface See Also
processes such as tectonics and impact cratering,
although no specific geological feature has been ▶ Albedo
identified at the surface of Ceres so far (besides ▶ Albedo Feature
some bright and dark spots that move with Ceres’ ▶ Ammonia
rotation) (Thomas et al. 2005; Li et al. 2006). ▶ Asteroid
Ceres is classified as a C- or G-type asteroid, ▶ Asteroid Belt, Main
sharing similarities with carbonaceous chon- ▶ Carbonaceous Chondrite
drites. Microwave dielectric measurements sug- ▶ Carbonate, Extraterrestrial
gest that Ceres is covered with dry claylike ▶ C-Asteroid
material at least 3 cm thick (Webster ▶ Clay
et al. 1988). Thermal emission spectroscopy ▶ Core, Planetary
also indicates the presence of iron-poor olivine, ▶ Crater, Impact
implying the presence of dry silicate, possibly on ▶ Crust
top of phyllosilicates (Witteborn et al. 2000). ▶ Cryovolcanism
Spectral signatures of ▶ carbonates and iron- ▶ Differentiation, Planetary
rich phyllosilicates have also been identified on ▶ Dwarf Planet
the surface of Ceres and are distributed globally ▶ Heat Flow, Planetary
in constant amount throughout the surface ▶ Heat Transfer, Planetary
(Rivkin et al. 2006). A 3–4 mm spectral feature ▶ Hydrosphere
is attributed to ▶ ammonia-bearing clay, either ▶ Hydrostatic Equilibrium
CH 423
▶ Hydrothermal Environments Witteborn FC, Roush TL, Cohen M (2000) Thermal emis-
▶ Interior Structure, Planetary sion spectroscopy of 1 Ceres: evidence for olivine,
thermal emission spectroscopy and analysis of dust,
▶ Mantle disks, and regoliths. In: Sitko ML, Sprague AL, Lynch
▶ Minor Planet DK (eds) Proceedings of a meeting held at LPI April
▶ Phyllosilicates, Extraterrestrial 1999: Astronomical Society of the Pacific Conference
▶ Primordial Heat Series, vol 196, pp 197–203
▶ Regolith, Planetary C
▶ Rock
▶ Rotation Planet
Cerium, Anomalies of
▶ Silicate Minerals
▶ Space Weathering
Francis Albarède
▶ Sun (and Young Sun)
Ecole Normale Supérieure de Lyon,
▶ Water
Lyon, France
▶ Water Activity
Definition
Cerium (Ce) is the second rare-earth element
Castillo-Rogez JC, McCord TB (2009) Ceres’ evolution
and present state constrained by shape data. Icarus (REE) in the periodic table, most of which are
203:443–459. doi:10.1016/j.icarus.2009.04.008 trivalent at the conditions prevalent in the mantle,
King TVV, Clark RN, Calvin WM, Sherman DM, Brown the crust, and planetary surfaces. In the modern
RH (1992) Evidence for ammonium-bearing minerals ocean, the redox boundary between trivalent and
on ceres. Science 255:1551–1553
Li J-Y, McFadden LA, JWm P, Young EF, Stern SA, tetravalent Ce is close enough to the O2H2O
Thomas PC, Russell CT, Sykes MV (2006) Photomet- equilibrium that a substantial fraction of this ele-
ric analysis of 1 Ceres and surface mapping from HST ment is oxidized and rapidly scavenged by Fe and
observations. Icarus 182:143–160 Mn oxides. Excess Ce with respect to adjacent
McCord TB, Sotin C (2005) Ceres: evolution and current
state. J Geophys Res 110:E05009 REE La and Pr is common in Mn nodules and
McCord TB, Castillo-Rogez JC, Rivkin A (2011) Ceres: encrustations, which leaves seawater, phos-
its origin, evolution and structure and Dawn’s potential phates, and carbonates with a deficit. Cerium
contribution. Space Sci Rev 163:63–76 anomalies in sediments and sedimentary rocks
Rivkin AS, Volquardsen EL, Clark BE (2006) The surface
composition of Ceres: discovery of carbonates and are useful proxies for the state of oxidation in
iron-rich clays. Icarus 185:563–567 the ocean and the atmosphere in the past.
Rivkin AS, Li JY, Miliken RE, Lim LF, Lovell AJ, Schmidt
BE, McFadden LA, Cohen BA (2011) The surface
composition of Ceres. Space Sci Rev 163:95–116
Russell CT, Raymond CA (2011) The Dawn mission to See Also
Vesta and Ceres. Space Sci Rev 163:3–23
Russell CT, Capaccioni F, Coradini A, de Sanctis MC, ▶ Chert
Feldman WC, Jaumann R, Keller HU, McCord TB, ▶ Great Oxygenation Event
McFadden LA, Mottola S, Pieters CM, Prettyman TH,
Raymond CA, Sykes MV, Smith DE, Zuber MT ▶ Ocean, Chemical Evolution of
(2007) Dawn mission to Vesta and Ceres. Earth ▶ Oxygenation of the Earth’s Atmosphere
Moon Planet 101:65–91
Thomas PC, JWm P, McFadden LA, Russell CT, Stern SA,
Sykes MV, Young EF (2005) Differentiation of the aster-
oid ceres as revealed by its shape. Nature 437:224–226
Webster WJ, Johnston KJ, Hobbs RW, Lamphear ES,
Wade CM, Lowman PD, Kaplan GH, Seidelmann PK
CH
(1988) The microwave spectrum of asteroid ceres.
Astron J 95:1263–1268. doi:10.1086/114722 ▶ Methylidyne
424 CH+
CH+ CH3CHNH
▶ Methylidyne Cation ▶ Ethanimine
CH2 CH3CN
▶ Methylene ▶ Acetonitrile
CH2CHCN CH3COOCH3
▶ Vinyl Cyanide ▶ Methyl Acetate
CH3 CH3O
▶ Methyl Radical ▶ Methoxy Radical
CH3C6H CH3OCH3
▶ Methyl Triacetylene ▶ Dimethyl Ether
CH3CH2CHO CH3SH
▶ Propionaldehyde ▶ Methanethiol
CH3CH2CN CH4
▶ Ethyl Cyanide ▶ Methane
CH3CHCH2 Chalcedony
▶ Propylene ▶ Chert
Chance and Randomness 425
Synonyms
Chalcophile Elements
Haphazardness; Indeterminacy; Stochasticity;
Francis Albarède Uncertainty
France
Definition C
Chance and randomness are usually considered

Definition as synonymous; however, they can have different
meanings, in several scientific fields as in every-
In the Berzelius-Goldschmidt classification, day contexts. In particular, chance has a broader
chalcophile elements are elements with a low scope than randomness, the latter being often
affinity for oxygen and which preferentially interpreted according to more specific mathemat-
bond with sulfur to form sulfides. Their name ical connotations. Broadly speaking, both are
derives not from sulfur but from copper, which used to qualify events that are unpredictable in
also forms sulfides. This group comprises transi- the sense that they have no particular aim or
tion elements (Cu, Zn, Cd, Ag, Hg), heavy metals direction (unbiased events) and that they occur
(Ga, In, Sn, Pb, Po, Bi, Tl), and metalloids (Ge, S, in an irregular and disordered (haphazard) way
Sb, Se, Te, As). As a consequence of their rela- which makes it difficult to make money betting.
tively low condensation temperatures
(500–1,100 K), most of these elements are
depleted in terrestrial planets with respect to Overview
chondrites.
Chance is a double-faced notion including subjec-
tive chance, which concerns our knowledge of real
events, and objective chance, which refers to an
See Also inherent property of the structure of the world,
independently of our knowledge. Chance and ran-
▶ Lithophile Elements domness are often used as counterparts to deter-
▶ Siderophile Elements minism: a random process is said to be
nondeterministic in the sense that, from a set of
starting conditions, it can produce different out-
comes according to some law of probability. How-
ever, neither notion is necessarily incompatible
Chance and Randomness with the assumption of determinism. Subjective
chance, when defined as ignorance of the real
Francesca Merlin underlying causes, implies that there is no chance
CNRS UMR 8690 IHPST & Université Paris 1, in the real world, but for human knowledge: for
Paris, France instance, we might assign a 50 % chance to both
possible outcomes of the flip of a fair coin
(“heads” and “tails”) because we ignore the under-
Keywords lying causes (e.g., the way the coin is flipped). This
recalls the Laplacian notion of chance, which is a
Chance; Determinism; Indeterminism; Probabil- deterministic notion; nevertheless, subjective
ity; Randomness; Unpredictability chance is also compatible with indeterminism in
426 Chandrasekhar’s Limit
so far as the underlying causes that we do not Hacking I (1995) The taming of chance. Cambridge Uni-
know about might still be the result of an indeter- versity Press, Cambridge, 1990
Heisenberg W (1958) Physics and philosophy. Harper,
ministic process. Objective chance can refer to an New York
event that is not planned (or by design), as in Laplace P-S (1814) A philosophical essay on probabilities
Aristotle’s accidental meeting with the person (English edition: Laplace P-S 1951). Dover, New York
who owed him money. This notion implies the Martin-Löf P (1966) The definition of random sequences.
Inf Control 9:602–619
confluence of two or more independent causal Poincaré H (1921) The foundations of science: science and
chains: in this sense, Cournot claimed that the hypothesis, the value of science, science and method
fact that two brothers serving in different armies (trans: Halsted GB). The Science Press, New York
died the same day is a matter of chance. Chaotic
processes provide an example of chance as sensi-
tivity to initial conditions in the sense that small
Chandrasekhar’s Limit
differences in initial conditions may yield radi-
cally different outcomes. Whereas all notions of
Daniel Rouan
chance mentioned above are noncommittal to
determinism nor indeterminism, this is not the
Meudon, France
case for chance according to the Copenhagen
interpretation of quantum mechanics, according
to which indeterminism is considered a true
Definition
description of the microlevel world. Randomness
mostly has a mathematical connotation, like in
Chandrasekhar’s limit, named after the Indian
algorithmic information theory where a binary
astrophysicist Subrahmanyan Chandrasekhar, is
sequence is said to be Kolmogorov random if
a critical mass of about 1.4 solar masses that the
and only if it is incompressible, i.e., it is shorter
core of a ▶ star can attain, before collapsing to
than any computer program that can produce
become a neutron star or a black hole. This core is
it. Other formal definitions of a random sequence
built from the heavy elements that the star syn-
(e.g., Martin-Löf’s) have been recently formu-
thesizes during its lifetime through nuclear
lated, each of which tries to capture our intuitive
fusion. This limit is thus the maximum mass of
notion of randomness.
a stable ▶ white dwarf star. Beyond this critical
mass, the relativistic electron degeneracy pres-
sure is unable to counteract the gravitational
See Also forces. Stars with mass higher than eight solar
masses develop a degenerate core, whose mass
▶ Materialism will grow until it exceeds this limit, leading to a
▶ Physicalism ▶ supernova explosion.
▶ Reductionism
▶ Vitalism
See Also
▶ Supernova
Aristotle (1984) Physics. In: Barnes J (ed) The complete ▶ White Dwarf
works of Aristotle, vol I and II. Princeton University
Press, Princeton
Calude CS (ed) (2007) Randomness and complexity from
Leibniz to Chaitin. World Scientific, Singapore
Cournot AA (1843) Exposition de la Théorie des Chances
et des Probabilités. Hachette, Paris
Channels
Earman J (1986) A primer on determinism. D. Reidel
Publishing, Dordrecht ▶ Valley Networks
Charge Transfer 427
Chaotic Region Characterization of Microfossils
Daniela Tirsch ▶ Microfossils, Analytical Techniques

Charge Exchange C
Synonyms
▶ Charge Transfer
Chaotic terrains
Definition Charge Transfer
A chaotic region is a distinctive area of fractured Steven B. Charnley

and disintegrated terrain. It is characterized by a Solar System Exploration Division, Code 691,
textured matrix featuring randomly orientated Astrochemistry Laboratory, NASA Goddard
flat-topped mesas, knobs of various sizes, and Space Flight Center, Greenbelt, MD, USA
deep depressions. Chaoses are primarily found
on ▶ Mars in regions unofficially called “chaotic
terrains.” The latter is located east of ▶ Valles Keywords
Marineris and involves parts of the huge ▶ out-
flow channel floors. Various processes are Chemical reactions; Interstellar medium
suggested to have formed the Martian chaoses,
including disruption and collapse of an icy ▶ per-
mafrost layer initiated by pressurized groundwa- Synonyms
ter, the action of aqueous processes, and multiple
episodes of nested surface collapses caused by Charge exchange
the removal of subsurface material, either ground
ice or magma. Chaos regions can also be found on
▶ Jupiter’s moon ▶ Europa, which is suggested Definition
to be sites of melt-through from the subsurface.
Charge transfer is a chemical process whereby
electric charge is transferred from a positive or
See Also negative ion (cation or ▶ anion) to a neutral atom
or molecule.
▶ Europa
▶ Jupiter
▶ Mars Overview
▶ Outflow Channels
▶ Permafrost Simple charge transfer reactions between atomic
▶ Valles Marineris or molecular ions and neutral atoms and mole-
cules are important in many astronomical envi-
ronments, ranging from ▶ planetary nebulae to
▶ comets, to dark ▶ molecular clouds. Reactions
Chaotic Terrains of the type
▶ Chaotic Region Hþ þ H2 O ! H2 Oþ þ H
428 Charge-Coupled Device
of 19,640 km or 17 Pluto radii. The orbital period

and
of the Pluto-Charon system is 6.4 days. Its diam-
eter is 1,206 km and its density is 1.8 g/cm3. With
C2 H2 þ CN ! CN þ C2 H2
a mass ratio of about 9, both Pluto and Charon can
be considered as parts of a binary Pluto-Charon
are ▶ exothermic, rapid, and proceed with no
system. Charon’s properties have been studied
molecular rearrangement. On the other hand, disso-
through photometry during mutual eclipses and
ciative charge transfer reactions involve the break-
transits and stellar occultations. Unlike Pluto’s,
ing of chemical bonds, examples of which are:
the surface of Charon is mostly made of water ice
with no evidence of atmosphere. Numerical sim-
Heþ þ CO ! Cþ þ O þ He
ulations indicate that Charon was formed after a
giant impact that took place some 4.5 Gy ago, in a
and
scenario comparable to the Earth-Moon
formation.
C2 H þ N ! CN þ CH:
See Also
See Also
▶ Anion
▶ Comet ▶ Kuiper Belt
▶ Exothermic ▶ Pluto
▶ Interstellar Chemical Processes ▶ Trans-Neptunian Object
▶ Molecular Cloud

Chasma, Chasmata
Millar TJ, Williams DA (eds) (1988) Rate coefficients in
astrochemistry. Kluwer, Dordrecht
Roland J. Wagner
Charge-Coupled Device
▶ CCD Synonyms
Canyon
Charon
Therese Encrenaz Definition

Meudon, Meudon, France A chasma (plural: chasmata) is a broad, deep,
elongated trough or depression. Chasmata are
bounded by steep scarps that can form a series
Definition of terraces. A chasma is preferentially created by
extensional tectonic forces. The ▶ terrestrial
Charon, discovered in 1978 by James Walter planets, ▶ Venus and ▶ Mars, have a large num-
Christy, is orbiting around ▶ Pluto at a distance ber of chasmata on their surface. In the outer
Chassignites 429
▶ Solar System, chasmata are major surface fea- The crystallization ages of the samples have
tures on the icy satellites of ▶ Saturn and been determined to lie around 1.3 Ga,
▶ Uranus. corresponding to middle Amazonian epoch
(Bogard and Garrison 1999; Nakamura
et al. 1982; Nyquist et al. 2001), similar to the
nakhlites crystallization age. The similarity with
See Also C
the crystallization history of the nakhlites and a
nearly identical initial 87Sr/86Sr isotopic ratio led
▶ Fossa, Fossae
first to the conclusion that both the chassignites
▶ Mars
and the nakhlites may sample different sites of
▶ Rima, Rimae
the same or similar igneous complexes
▶ Rupes, Rupēs
(Nakamura et al. 1982). Moreover, the similar
▶ Satellite or Moon
cosmic-ray exposure age (11.1 1.6 Ma) implies
▶ Saturn
that they all have been ejected in one event
▶ Sulcus, Sulci
(Nyquist et al. 2001). However, their REE abun-
▶ Terrestrial Planet
dances indicate that they have not crystallized
▶ Uranus
from the same magma (Longhi 1991; Wadhwa
▶ Venus
and Crozaz 1994). Harvey and Hamilton (2005)
proposed the northeast region of Syrtis Major
volcanic complex to be a possible site for the
ejection of chassignites. Although some sites,
Chassignites e.g., Nili Fossae, that were proposed as source
terrains for chassignites are of Noachian age, the
Ana-Catalina Plesa olivine-bearing regions within these terrains
German Aerospace Center (DLR), Institute of could be much younger.
Keywords See Also
SNC meteorites; Isotopic ratio; Cosmic-ray ▶ Isotopic Fractionation (Planetary Process)

exposure age; Noachian; Amazonian ▶ Isotopic Ratio
▶ Crater, Impact
▶ Meteorites
Definition
Chassignites are a subgroup of the SNC meteor- References and Further Reading
ites named after the first sample found, the
Bogard DD, Garrison DH (1999) Argon-39-argon-40 ages
Chassigny meteorite. These dunitic samples and trapped argon in Martian Shergottites, Chassigny,
(more than 90 % olivine) are classified as and Allan Hills 84001. Meteorit Planet Sci 34:451–473
olivine-chromite cumulate rocks. Harvey RP, Hamilton EV (2005) Syrtis major as the
source region of the nakhlite/Chassigny group of Mar-
tian meteorites: implications for the geological history
of Mars. Proc Lunar Planet Sci Conf 36:1019
Overview Longhi J (1991) Complex magmatic processes on mars:
inferences from the SNC meteorites. Proc Lunar Planet
Sci 21:695–709
This subgroup of the Martian meteorites contains
Nakamura N, Kogi H, Kagami H (1982) Rb-Sr isotopic
two samples: Chassigny and NWA 2737 (field and REE abundances in the Chassigny meteorite.
name “Diderot”) sample. Meteoritics 17:257–258
430 Chassigny
Nyquist LE, Bogard DD, Shih C-Y, Greshake A, Stöer D,

Eugster O (2001) Ages and geologic histories of Mar- Chemical Bistability
tian meteorites. Chronol Evol Mars 96:105–164
Wadhwa M, Crozaz G (1994) Rare earth element distri-
butions in Chassigny: clues to its petrogenesis and Steven B. Charnley
relation to the nakhlites. Proc Lunar Planet Sci Conf Solar System Exploration Division, Code 691,
25:1451–1452 Astrochemistry Laboratory, NASA Goddard
Chassigny
Definition
Ana-Catalina Plesa
Chemical bistability in astrochemistry refers to
the existence of multiple steady states known to
occur in chemical models of molecular abun-
dances in dense interstellar clouds. For certain
combinations of model (control) parameters
Definition
(cosmic ray ionization rate, elemental abun-
dances, etc.), three steady states can appear, com-
Chassigny is a 4-kg Martian meteorite found in
prising two stable states connected by an unstable
1815 in Chassigny, France. It names the SNC sub-
one, i.e., the solutions exhibit hysteresis.
group containing also the NWA 2737 (Diderot)
sample. Chassigny is a dunitic sample (more than
90 % olivine), classified as an olivine-chromite
cumulate rock, for which a crystallization age has
History
been determined to lie around 1.36 Ga, similar to
the nakhlites age. Spectral signatures matching the
Chemical bistability in astrochemical models was
mineralogy of Chassigny have been identified in
first positively identified by Le Bourlot
Nili Fossae near Syrtis Major, southwestern region
et al. (1993).
of Isidis rim, in Ganges and Eos Chasmata, within
craters southwest of Ganges Chasma, and in the
northeastern part of the Hellas basin rim. Although
some candidate sites like Nili Fossae have an old
See Also
Noachian age, the olivine-bearing materials
located within could be much younger.
▶ Molecular Cloud
See Also
▶ Crater, Impact References and Further Reading

▶ Isotopic Fractionation (Planetary Process)
Le Bourlot J, Pineau des Forets G, Roueff E, Schilke
▶ Isotopic Ratio P (1993) Bistability in dark cloud chemistry.
▶ Meteorites Astrophys J 416:L87–L91
Chemical Adsorption Chemical Evolution
▶ Chemisorption ▶ Prebiotic Chemistry

Chemical Reaction Network 431
A reaction network is typically described by

Chemical Fossil its stoichiometric matrix n. The ni,j element is the
stoichiometric coefficient of the compound i in
▶ Biomarkers the reaction j, i.e., the number of molecule of
i that is involved in the reaction j. This number
is by convention negative for the reactants (that
are disappearing) and positive for the products C
Chemical Reaction Network (that are formed). The mathematical analysis of
this matrix gives information about the structure
Raphaël Plasson of the reaction network (Schilling et al. 2000).
Department of Earth and Planetary Science, The identification of matter fluxes and of patterns
Harvard University, Cambridge, MA, USA of mass conservation leads to the description of
the reaction network in terms of transformation
pathways (Papin et al. 2004) and of conserved
Keywords moieties (Schuster and Hilgetag 1995). This
mathematical analysis only gives static informa-
Metabolism; Stoichiometric matrix tion about the reaction network. Adding the
kinetic data relative to each chemical reaction
allows establishment of the set of ordinary differ-
Synonyms ential equations (ODE) describing the reaction
network. Its numerical integration gives the
Automaton, Chemical; Chemical system; Reac- dynamical behavior of the system (Alves
tion network et al. 2006).
The description of chemical reaction networks
is used in very different fields, dealing with
Definition chemical complexity. This is typically the case
of biosystems, especially for the description of
A chemical reaction network is a set of chemical ▶ metabolism (Schuster and Hilgetag 1995;
compounds (being the network nodes) connected Schilling et al. 2000; Papin et al. 2004; Metabo-
by a set of chemical reactions (being the network lism databases), but also in abiotic systems as in
vertices). astrochemistry (Woodall et al. 2007), combustion
modeling (Manion et al. 2008), etc. The precise
description of these complex chemical systems
Overview generally relies on the existence of extensive
databases, summing up thermodynamic and
A chemical system is a set of chemical compounds kinetic data.
that are involved in a set of chemical reactions. In the field of astrobiology, ▶ prebiotic chem-
Formally, this can be described as a network, istry can be seen as a bridge between ▶ abiotic
where each compounds are connected with each and biotic chemical reaction networks. The pur-
other by the corresponding chemical transforma- pose is to understand how a reaction network
tions; each chemical compound is then a node of consisting of very simple compounds can spon-
the network, while each chemical reaction is a taneously evolve into a more complex and struc-
directed vertex of the network, connecting the tured network, and how complex behaviors can
involved chemical compounds, from the reactants emerge from the association of several simple
towards the products. The purpose is to study this reactions (e.g., network ▶ autocatalysis, energy
system as a whole, rather than solely considering it coupling, self-replication, etc.) (Wagner and
as the sum of its different elements. Ashkenasy 2009).
432 Chemical System
See also
Chemical Zones
▶ Automaton, Chemical
▶ Chemical Evolution ▶ Redox Zonation
▶ Cosmochemistry
▶ Metabolism
Chemiosmotic Potential
▶ Proton Motive Force

Alves R, Antunes F, Salvador A (2006) Tools for kinetic

modeling of biochemical networks. Nat Biotechnol
24:667–672
Manion JA, Huie RE, Levin RD, Burgess DR Jr, Orkin Chemisorption
VL, Tsang W, McGivern WS, Hudgens JW, Knyazev
VD, Atkinson DB, Chai E, Tereza AM, Lin CY, Steven B. Charnley
Allison TC, Mallard WG, Westley F, Herron JT,
Hampson RF, Frizzell DH (2008) NIST chemical
kinetics database. National Institute of Standards and 691, Astrochemistry Laboratory, NASA Goddard
Technology, Gaithersburg, http://kinetics.nist.gov/ Space Flight Center, Greenbelt, MD, USA
Metabolism databases: KEGG, http://www.genome.ad.jp/
kegg/reaction/; Biocyc, http://biocyc.org/; UniPathway,
http://www.grenoble.prabi.fr/obiwarehouse/unipathway
Papin JA, Stelling J, Price ND, Klamt S, Schuster S, Synonyms
Palsson BO (2004) Comparison of network-based
pathway analysis methods. Trends Biotechnol Chemical adsorption
22:400–405
Schilling C, Letscher D, Palsson BO (2000) Theory for
the systemic denition of metabolic pathways and
their use in interpreting metabolic function from Definition
a pathway-oriented perspective. J Theor Biol
203:229–248
Schuster S, Hilgetag C (1995) What information about the
Chemisorption involves the formation of cova-
conserved-moiety structure of chemical reaction sys- lent chemical bonds between an adsorbed atom or
tems can be derived from their stoichiometry? J Phys molecule (or its dissociation products) and the
Chem 99:8017–8023 molecules present in a solid surface, such as that
Wagner N, Ashkenasy G (2009) Symmetry and order in
of an interstellar dust grain. This strong bonding
systems chemistry. J Chem Phys 130:164907
Woodall J, Agúndez M, Markwick-Kemper AJ, Millar TJ contrasts with the weaker bonding during
(2007) The UMIST database for astrochemistry 2006. ▶ physisorption.
Astron Astrophys 466:1197–1204
See Also
Chemical System ▶ Adsorption

▶ Chemical Reaction Network ▶ Physisorption
Chemocline 433
▶ Energy Conservation
Chemoautotroph ▶ Energy Sources
Ricardo Amils ▶ Extremophiles
Departamento de Biologia Molecular, ▶ Hot Vent Microbiology
Universidad Autónoma de Madrid, Madrid, ▶ Iron Cycle
Spain ▶ Lithotroph C
▶ Prokaryote
▶ Respiration
Definition
Chemoautotrophs are organisms that obtain their

energy from a chemical reaction (chemotrophs)
but their source of carbon is the most oxidized Chemocline
form of carbon, carbon dioxide (CO2). The best
known chemoautotrophs are the chemolithoau- Daniele L. Pinti
totrophs that use inorganic ▶ energy sources, GEOTOP Research Center for Geochemistry and
such as ferrous iron, hydrogen, hydrogen sulfide, Geodynamics, Université du Québec à Montréal,
elemental sulfur or ammonia, and CO2 as their Montréal, QC, Canada
▶ carbon source. All known chemoautotrophs are
prokaryotes, belonging to the ▶ Archaea or
▶ Bacteria domains. They have been isolated in Definition
different extreme habitats, associated to deep-sea
vents, the deep biosphere, or acidic environ- The chemocline is the vertical chemical gradient
ments. This form of ▶ energy conservation is (cline) which is produced in a mass of water
considered one of the oldest on Earth. These having restricted circulation (for example, deep
microorganisms are of astrobiological interest lakes). The local conditions favor the formation
because they could develop in the extreme con- of bottom anoxic waters, where only anaerobic
ditions existing in different extraterrestrial plan- forms of life exist, overlain by oxygenated
etary bodies, like Mars or Europa. waters, where aerobic photosynthetic life flour-
ishes. At the contact between these two zones,
photosynthetic communities of anaerobic purple
See Also sulfur bacteria develop, taking advantage of both
the sunlight from above and the hydrogen sulfide
▶ Acidophile (H2S) produced below. In astrobiology, this zone
▶ Autotroph of the chemocline is studied because it could be
▶ Autotrophy an analog of Archean niches of life where limited
▶ Bacteria oxygenated waters developed over an anoxic
▶ Bioenergetics Archean ocean.
▶ Carbon Source
▶ Chemotroph See Also
▶ Deep-Sea Microbiology
▶ Deep Subsurface Microbiology ▶ Photosynthesis
▶ Early Archean ▶ Redox Zones
434 Chemolithoautotroph
▶ Carbon Dioxide
Chemolithoautotroph ▶ Chemotroph
▶ Electron Transport
Ricardo Amils ▶ Nitrification
Centro de Biologia Molecular, Universidad ▶ Proton Motive Force
Keywords Chemolithotroph
Hydrogen oxidizers; Iron oxidizers; Nitrifying Ricardo Amils

bacteria; Sulfur oxidizers Departamento de Biologia Molecular,
Universidad Autónoma de Madrid, Madrid,
Spain
Definition
A chemolithoautotroph is an autotrophic microor- Keywords

ganism that obtains energy by oxidizing inorganic
compounds. Most ▶ chemolithotrophs are ▶ auto- Chemolithotrophy
trophs. Examples of relevant inorganic electron
donors include hydrogen, hydrogen sulfide, fer-
rous iron, and ammonia. Winogradsky described Definition
the concept of chemolithoautotrophy for the first
time while studying the ammonia-oxidizing bac- A chemolithotroph is an organism that is able to
teria. Chemolithoautotrophic organisms have use inorganic reduced compounds as a source of
▶ electron transport complexes, similar to those energy. This mode of metabolism is known as
of chemoorganotrophs, which are used to generate chemolithotrophy.
a ▶ protein motive force. The proton motive force
drives the synthesis of ATP. In this case, the
reduction of CO2 requires the use of ATP and
reducing power, which is, most often, obtained History
through the use of the electron transport chain in
reverse mode, consuming energy. Sulfur, iron, and Chemolithotrophy was discovered by
ammonia oxidizers are fundamental elements in Winogradsky while studying the microorganisms
the biogeochemical cycles of these elements. involved in the oxidation of sulfur compounds.
Chemolithoautotrophic organisms are of special
interest to astrobiology due to their minimal
requirements for development and their ability to
readily adapt to extreme conditions. Overview
Chemolithotrophy is found only in prokaryotes

See Also and is widely distributed among Bacteria and
Archaea. The spectrum of inorganic compounds
▶ ATP Synthase that can be used as electron donors by
▶ Bioenergetics chemolithotrophs is rather broad (H2S, S0,
Chemolithotroph 435
S2O32, H2, Fe2+, NO2or NH3). Some microor- elemental sulfur (S0), and thiosulfate (S2O32).
ganisms are rather specific regarding the inor- The final product of sulfur oxidation is sulfate
ganic substrates they can use to generate energy, (SO42), although an intermediate step is the
while others are able to use different compounds formation of elemental sulfur, which in some
(versatile). The best characterized cases is stored as an alternative source of energy.
chemolithotrophs are aerobic respirers, which One of the products of sulfur oxidation reaction is
use oxygen as the ▶ electron acceptor, although the generation of protons (H+), consequently one C
the list of chemolithotrophs capable of employing result of the oxidation of reduced sulfur com-
▶ anaerobic respiration is increasing rapidly. pounds is the acidification of the environment
Chemolithotrophs have ▶ electron transport sys- by the production of sulfuric acid.
tems similar to those of chemoorganotrophs, The aerobic oxidation of ferrous iron (Fe2+) to
which are used for the generation of a ▶ proton ferric ▶ iron (Fe3+) is an energy-yielding reac-
motive force. The only difference is that tion, used by some prokaryotes to conserve
chemolithotrophs donate electrons directly to energy. Only a small amount of energy is gener-
the electron transport chain, while chemoorga- ated by this reaction, thus iron-oxidizing micro-
notrophs must generate cellular reducing power organisms must oxidize large amounts of reduced
(NADH) from the ▶ oxidation of reduced organic iron to grow. Ferrous iron is oxidized very rapidly
compounds, which is then used to donate elec- in the presence of oxygen, while it is very stable
trons to the electron transport system. This proton at acidic conditions. This is probably the reason
motive force is used to generate ATP or any why many iron-oxidizing microorganisms are
cellular functions that might require this type of acidophilic. Despite the instability of ferrous
energy (active transport, movement, etc.). An iron at neutral pH, there are a number of iron-
important distinction between chemolithotrophs oxidizing bacteria that can thrive at
and chemoorganotrophs is their source of carbon. circumneutral pH. Some anoxigenic phototrophic
Chemoorganotrophs use organic compounds as bacteria can use ferrous iron as a source of envi-
both energy and carbon sources, while ronmental reducing power. Recently, it has been
chemolithotrophs are generally autotrophs (with shown that some denitrifying bacteria can anaer-
few exceptions, known as mixotrophs, that use obically respire (oxidize) reduced iron. The use
reduced organic compounds as a source of car- of ferrous iron to obtain energy is widely distrib-
bon). Chemolithotrophs can obtain the reducing uted in nature, a property that was ignored until
power needed to assimilate CO2 directly from the recently, due to thermodynamic considerations.
inorganic substrate (only H2 oxidizers) or by the The most common nitrogen compounds used
reverse electron transport reaction (the rest of as electron donors for energy conservation are
chemolithotrophs), in this case using proton ammonia (NH3) and nitrite (NO2). Both com-
motive force as a source of energy. pounds can be oxidized aerobically by
Hydrogen is a common product of geochemi- chemolithotrophic nitrifying bacteria. Some
cal reactions and microbial metabolism, and a nitrifying microorganisms oxidize ammonia to
number of chemolithotrophs are able to use it as nitrite, while another group oxidizes nitrite to
an electron donor in energy metabolism. A wide nitrate. The complete oxidation of ammonia
variety of anaerobic H2-oxidizing Bacteria and requires the concerted activity of these two
Archaea are known, differing in the electron types of microorganisms. A special case of
acceptor they use (nitrate, sulfate, ferric iron, nitrogen-oxidizing microorganisms corresponds
etc.). to those capable of carrying out the anoxic oxi-
The most common sulfur compounds used as dation of ammonia, a process known as anamox.
electron donors are hydrogen sulfide (H2S), In this case, the electron acceptor is nitrite, and
436 Chemoorganotroph
the product of the metabolic reaction in addition González-Toril E, Gómez F, Malki M, Amils R (2006)
to proton motive force is the generation of N2. Isolation and study of acidophilic microorganisms. In:
Rainey F, Oren A (eds) Methods in microbiology,
This metabolic reaction is carried out by a special vol 35. Elsevier, Oxford, pp 463–502
type of microorganisms belonging to the Leininger S, Urich T, Schloter M, Schwark L, Qi J, Nicol
Planctomycetes phylum of Bacteria. GW, Prosser JI, Schuster SC, Schleper C (2006)
Due to their metabolic properties, Archaea predominate among ammonia-oxidizing pro-
karyotes in soils. Nature 442:806–809
chemolithotrophs are of astrobiological interest Madigan MT, Martinko JM, Dunlap PV, Clark DP
and also critical elements of the ▶ biogeochemi- (2009) Brock biology of microorganisms, 12th edn.
cal cycles. Cummings, Benjamin
Strous M, Fuerst JA, Kramer EH et al (1999) Missing
lithotroph identified as new planctomycete. Nature
400(6743):446–449. doi:10.1038/22749
See Also Winogradsky S (1949) Microbiology du Sol. Masson,
Parı́s
▶ Acidophile Yamanaka T (2008) Chemolithoautotrophic bacteria. Bio-
chemistry and environmental biology. Springer,
▶ Aerobic Respiration Tokyo
▶ ATP Synthase
▶ Autotrophy
▶ Bioenergetics Chemoorganotroph
▶ Biogeochemical Cycles
▶ Chemoautotroph Ricardo Amils
▶ Chemolithoautotroph Departamento de Biologia Molecular,
▶ Chemoorganotroph Universidad Autónoma de Madrid, Madrid,
▶ Electrochemical Potential Spain
▶ Electron Acceptor
▶ Electron Carrier
▶ Electron Donor Keywords
▶ Electron Transport
▶ Energy Conservation Fermentation; Heterotrophs; Respiration
▶ Energy Sources
▶ Iron
▶ Iron Cycle Definition
▶ NADH, NADPH
▶ Nitrogen Cycle, Biological A chemoorganotroph is an organism that obtains
▶ Oxidation energy from the oxidation of reduced organic
▶ Proton Motive Force compounds. The list of compounds from which
▶ Sulfur Cycle chemoorganotrophic organisms can generate
energy and their sources of carbon is very long,
making these microorganisms extremely versa-
tile. Two mechanisms for energy conservation
References and Further Reading are known for chemoorganotrophs: ▶ fermenta-
Ehrlich HL, Newman DK (2009) Geomicrobiology,
tion and ▶ respiration. In the case of fermenta-
5th edn. CRC Press, New York tion, cellular energy in the form of ATP is
Fernandez-Remolar DC, Morris RV, Gruener JE, obtained by cytoplasmatic soluble catalytic reac-
Amils R, Knoll AH (2005) The Rio Tinto basin, tions involved in substrate-level phosphorylation.
Spain: mineralogy, sedimentary geobiology, and
implications for interpretation of outcrop rocks at
In the case of respiration, ATP is produced at the
Meridiani Planum, Mars. Earth Planet Sci Lett expense of the ▶ proton motive force resulting
240(1):149–167. doi:10.1016/j.epsl.2005.09.043 from coupling the substrate oxidation reactions
Chemotaxis 437
via the generation of reducing power to the elec- causes the bacterium to tumble since each flagel-
tron transport chain. Due to their type of metab- lum rotates independently.
olism, chemoorganotrophs are fundamental Bacteria monitor chemical concentrations
elements of the carbon cycle. using multiple transmembrane receptors, named
methyl-accepting chemotaxis proteins (MCPs).
See Also Five MCP-mediating responses to specific attrac-
tant and repellent stimuli have been reported in C
▶ Aerobic Respiration E. coli: Tar, Tsr, Trg, Aer, and Tap. The signals
▶ ATP Synthase from these receptors are transmitted across the
▶ Catabolism cell membrane into the cytoplasm, where Che
▶ Chemotroph proteins (CheA, CheB, CheW, CheY, CheR,
▶ Energy Sources and CheZ) are activated.
▶ Proton Motive Force Signals from the receptors are received by the
CheA histidine kinase. CheA is coupled to trans-
membrane receptors (MCP) by the adaptor pro-
tein CheW. The activation of the receptor causes
Chemotaxis autophosphorylation of CheA that also phosphor-
ylates CheB and CheY. CheYP binds then to
Irma Marı́n the flagellar motor protein FliM, inducing a
Departamento de Biologı́a Molecular, change from CCW to CW rotation of flagella,
Universidad Autónoma de Madrid, Madrid, thus increasing the frequency of tumbles of the
Spain bacterium (Fig. 1).
A feedback mechanism that modulates the
methylation level of the MCP receptors controls
Keywords adaptation. Two enzymes, CheB and CheR, are
involved in this mechanism by interacting with
Motility; Two-component system the receptor and chemically modifying them.
CheB, activated by CheA, acts as a
methylesterase that removes methyls from the
Definition cytoplasmic part of the receptor. It works antag-
onistically with CheR, which is a
Chemotaxis is the process by which motile bac- methyltransferase. The higher the amount of
teria sense changes in their chemical environ- methylated residues of the receptor, the lower
ment and move to more favorable conditions. the sensitivity to stimuli. The result is an
enhancement of CheA autophosphorylation and,
thereby, transmission of a CW signal. When an
Overview attractant generates a signal, demethylation of the
receptor is induced, closing the feedback loop.
Bacteria, Archaea, and some eukaryotes use The system is continuously adjusted to envi-
two-component signaling pathways to detect ronmental chemical levels, remaining sensitive to
environmental conditions and bring about appro- small changes even under extreme chemical con-
priate changes in cellular behavior. This process centrations, since the system compares concen-
has been studied in Escherichia coli over the past trations along the movement path. When
40 years. necessary this is switched to go closer to or fur-
E. coli has several flagella per cell (4–19) that ther away from a higher concentration of attrac-
can rotate in two ways: counterclockwise (CCW) tant or a repellent, respectively. Other
that aligns the flagella causing the bacterium to mechanisms are involved in increasing the abso-
swim in a straight line and clockwise (CW) that lute value of the sensitivity on a given
438 Chemotroph
Chemotaxis, Fig. 1 Model of chemotaxis mechanism as described in the text. A, B, R, W, Y, and Z are CheA, CheB,
CheR, CheW, CheY, and CheZ proteins, respectively
background. E. coli, but not B. subtilis, possesses

the protein CheZ that enhances the rate of CheY Chemotroph
dephosphorylation.
Ricardo Amils
See Also Universidad Autónoma de Madrid, Madrid,
Spain
▶ Motility
Definition
References and Further Reading Chemotrophs are organisms that obtain energy by
the oxidation of reduced compounds. The sub-
Bren EM (2000) How signals are heard during bacterial
chemotaxis: protein-protein interactions in sensory strates used by chemotrophs can be organic
signal propagation. J Bacteriol 182:6865–6873 (organotrophs) or inorganic compounds
Madigan MT, Martinko JM, Dunlap PV, Brock DP (lithotrophs). According to the carbon source,
(2008) Biology of microorganisms, 12th edn. Clark
chemotrophs can be either chemoautotrophs or
Benjamin Cumming, San Francisco
Prescott HK (2007) Microbiology, 7th edn. Willey, chemoheterotrophs. Because chemoheterotrophs
McGraw-Hill Science, New York use reduced organic compounds as a source of
Cheops 439
energy and a source of carbon, they are usually detailed view of Cambrian life, yielding insights
called heterotrophs, although the term is mislead- into early animal evolution and the Cambrian
ing because, strictly, it only refers to the carbon explosion. The Chengjiang biota provides an
source. Chemoautotrophs use inorganic energy early Cambrian counterpart to the exceptionally
sources and are known as chemolithoautotrophs preserved biota of the middle Cambrian Burgess
or lithoautotrophs. Chemolithoheterotrophs are a Shale, Canada.
special kind of chemotroph that use inorganic C
compounds as an energy source and reduced
organic compounds as a carbon source. They
are known as mixotrophs. Chemotrophs use See Also
▶ fermentation and ▶ respiration to obtain
energy. Fermentation is restricted to ▶ Burgess Shale Biota
organotrophs. ▶ Cambrian Explosion
See Also
▶ Aerobic Respiration Cheops

▶ Autotrophy Michel Viso
▶ Bioenergetics CNES/DSP/SME, Vétérinaire/DVM, Astro/
▶ Carbon Dioxide Exobiology, Paris Cedex 1, France
▶ Chemoautotroph
▶ Chemolithotroph Keyword
▶ Chemoorganotroph
▶ Energy Conservation Exoplanet
▶ Fermentation
▶ Respiration
Definition
The CHaracterizing ExOPlanet Satellite

(CHEOPS) is a small mission selected by the
Chengjiang Biota, China European Space Agency (ESA) in October 2012
and then adopted in February 2014. It will be the
Thomas H. P. Harvey first mission dedicated to searching for ▶ transits
Department of Earth Sciences, University of by means of ultrahigh precision photometry on
Cambridge, Cambridge, UK bright stars already known to host planets. It will
provide the unique capability of determining
accurate radii for a subset of planets with a size
Definition ranging from Earth to Saturn which orbit bright
stars and for which the mass has already been
The Chengjiang biota is an assemblage of fossils estimated from ground-based spectroscopic sur-
first discovered in 1984 at Maotianshan, near veys. The knowledge of mass and diameter will
Chengjiang, Yunnan Province, China, and dated allow to derive the density and then to draw
to ca. 520 Ma, within the Cambrian Period of hypotheses about the composition of these
geologic time. Exceptional preservation of soft- planets. The 250 kg satellite will be equipped
bodied marine organisms provides a rare, with a 32 cm telescope. The payload is developed
440 Chert
under the responsibility of the University of Bern Belt in West Greenland (Nutman et al. 1984).
(Switzerland). The mission will be ready for The Isua cherts may have had a sedimentary
launch in 2017 in a Sun-synchronous orbit. origin or may represent volcanic rocks that were
hydrothermally altered and layered as a conse-
quence of intense plastic deformation (Nutman
See Also and Friend 2009; Polat and Frei 2005). Trace
elements indicate that the chert/▶ banded iron
formation units have seawater-like signatures
suggesting at least a component of sedimentary
origin as a marine chemical precipitate.
Chert In both the ▶ Pilbara (Australia) and the
▶ Barberton (South Africa) Greenstone Belts,
Tanja Elsa Zegers the oldest well-preserved volcanic sequences
Paleomagnetic Laboratory, Institute of Earth which contain stratiform chert units (Byerly
Sciences, Utrecht University, Utrecht, CD, et al. 1996; Van Kranendonk et al. 2002) formed
The Netherlands between ~3.57 and ~3.30 Ga. Despite the good
preservation of the greenstone sequence, the ori-
gin of the stratiform chert units is still controver-
Keywords sial. van den Boorn et al. (2007) distinguished
three different end members of silica derivation
Archean; Microfossils; Alteration on the basis of Si isotopic signatures: direct pre-
(hydrothermal); Sediments cipitation of silica from seawater on the seafloor,
alteration of a precursor rock (volcanic, carbon-
ate, siliciclastic) by addition of silica from sea-
Synonyms water, and silica precipitation from hydrothermal
vents. Only in the first case does the chemical and
Chalcedony; Flint; Jasper isotopic composition of chert reflect the chemis-
try and temperature of Archean seawater.
Chert is very resistant to surface weathering
Definition and associated chemical alteration. Therefore,
chert is often considered to be the ideal agent for
Chert is a microcrystalline rock consisting almost the preservation of very ancient and potentially
exclusively of silica. Chert may occur as strati- extraterrestrial biosignatures (Westall 2008).
form units, occasionally with laminar alternation
of colors (red, white, black, green), in dikes cross-
cutting the stratigraphy, or as nodules in carbon- Overview
ates. Some stratiform cherts in Archean
▶ greenstone belts are among the earliest sedi- Cherts, both sedimentary (authigenic) and as a
mentary rocks and preserve the oldest evidence secondary alteration product, form by precipita-
for life (▶ Apex Chert, ▶ Pilbara Craton, West- tion from aqueous fluids. The solubility of silica
ern Australia). (Dove and Rimstidt 1994; Fleming and Crerar
1982) in water is largely independent of its pH
for pH lower than 9 but increases exponentially
History for pH >9. Solubility of silica also increases with
increasing temperature and with increasing salin-
The oldest preserved chert units occur in the ity. During the Archean, precipitation of silica
highly metamorphic and deformed 3.6–3.7 Ga was most likely abiogenic and either derived
supracrustal units of the ▶ Isua Supracrustal originally from ocean-floor hydrothermal vents
Chert 441
or from regional precipitation from normal but carbonaceous and clastic sedimentary deposition
silica-supersaturated seawater. From ~1.8 Ga ago but are more likely to represent deposition of
(Maliva et al. 2005), the silica cycle in the oceans silica directly from oversaturated seawater. Silica
began to be governed, as it still is, by silica- is thought to have been released from nearby
secreting organisms. Radiolaria are thought to ocean-floor hydrothermal vents even though
have appeared in the late Cambrian diatoms in these can only rarely be documented. These
the Jurassic. Since then, most silica derived from cherts show no macroscopic biogenic textures, C
continental weathering and delivered by rivers to but have been studied extensively for
the oceans is biologically consumed (Laruelle microtextures that might be indicative of fossil-
et al. 2009). ized microorganisms and microbial mats (Schopf
For our understanding of the evolution and et al. 2002; Westall et al. 2006; Westall 2008;
conditions of early life, the chert units found in Sugitani et al 2013). Some of the most famous
early and mid-Archean stratigraphic sequences putative microtextures were found in the ▶ Apex
such as in the Pilbara Craton (Australia) and Chert (Schopf et al. 2002). The claim that those
Barberton Greenstone Belt (South Africa) are filamentous textures represented evidence for life
most important. Both sequences, the at 3.45 Ga was countered by studies showing that
Warrawoona Group in the Pilbara Craton and similar textures could be generated in hydrother-
the Onverwacht Group in the Barberton Green- mal systems by Fischer-Tropsch-type abiotic
stone Belt, consist predominantly of mafic to processes (Brasier et al. 2002).
ultramafic volcanic units, with minor felsic vol- Because chert is so resistant to erosion, it pro-
canic and associated volcanoclastic units vides a good record for any study of ▶ early Earth
(3.47–3.3 Ga). Both sequences are so similar conditions. Samples from the Marble Bar Chert
that they may have been part of a single terrain (▶ Pilbara, Australia) have been used for paleo-
at the time of deposition (Zegers et al. 1998). magnetic studies (Suganuma et al. 2006). Oxygen
Chert units (typically less than 20 m thick) cap and Si isotopic compositions in chert have been
volcanic sequences and can commonly be traced used to infer the temperature of the Archean
over large (>50 km) distances (Kato and ocean (Knauth and Lowe 2003; Robert and
Nakamura 2003). A few thicker chert units Chaussidon 2006) of up to 70 C. However, van
occur in both the Pilbara Craton and Barberton den Boorn et al. (2010) showed that the samples
Greenstone Belt: the Strelley Pool Chert and the used in those studies were most likely deposited
Buck Reef Chert (350 m, Hofmann and Harris from hot hydrothermal water rather than from
2008), respectively. The detailed study of the ambient seawater. They suggest a maximum tem-
Strelley Pool Chert (3.43 Ga) by Allwood perature of ambient seawater of 55 C.
et al. (2006) showed that the chert consists of at Because of its formation by precipitation
least four members with different characteristics from water and the potential to preserve textural
and compositions, some of which show excel- and geochemical proxies for fossil life, chert
lently preserved ▶ stromatolites. The unit was is one of the targets to detect former life on
most likely deposited as a dolomite/carbonate Mars (Westall 2008). The ▶ Mars Exploration
reef on a shallow platform consisting of felsic Rover Spirit recently found amorphous silica
volcanoclastic sediments. Tice and Lowe (2004) (a precursor to chert) in the Gusev Crater (Rice
studied in detail the Buck Reef Chert, and sedi- et al. 2010).
mentary structures were interpreted to represent
deposition in a shallow- to deep-marine environ-
ment, with organic matter in the shallow-water Key Research Findings
facies resulting from photosynthetic
microbial mats. Chert units in Archean greenstone belts represent
Other chert units in the earliest greenstone the oldest sedimentary units in otherwise entirely
sequences show no strong evidence of volcanic sequences. Stratiform chert units may be
442 Chert
the result of direct precipitation from ▶ Banded Iron Formation

oversaturated seawater or may represent hydro- ▶ Barberton Greenstone Belt
thermal deposits. In some cases, units that origi- ▶ Barberton Greenstone Belt, Sedimentology
nally contain largely carbonate (dolomite, ▶ Barberton Greenstone Belt, Traces of Early
siderite) have been pervasively silicified. Some Life
of those units contain stromatolites and other ▶ Biomarkers
micromorphological indications of biological ▶ Biomarkers, Morphological
activity. ▶ Carbonate
▶ Microfossils
Applications ▶ Pilbara Craton
▶ Precambrian Oceans, Temperature of
Chert is very resistant to weathering and chemi-
cal alteration under surface conditions. This
makes chert units, if they can be shown to have References and Further Reading
formed in the Archean, ideal recorders of
Archean conditions, to be probed by a variety of Allwood AC, Walter MR, Kamber BS, Marshall CP,
Burch IW (2006) Stromatolite reef from the early
techniques.
Archaean era of Australia. Nature 441(7094):714–718
Brasier MD, Green OR, Jeplhcoat AP, Kleppe AK, Van
Kranendonk MJ, Lindsay JF, Steele A, Grassineau NV
Future Directions (2002) Questioning the evidence for Earth’s oldest
fossils. Nature 416(6876):76
Byerly G, Kroner A, Lowe D, Todt W, Walsh M (1996)
Chert units will continue to be studied as some of Prolonged magmatism and time constraints for sedi-
the best recorders of the conditions of early life. ment deposition in the early Archean Barberton green-
To correctly interpret micromorphological and stone belt: evidence from the upper onverwacht and fig
tree groups. Precambrian Res 78(1–3):125–138
geochemical signatures, the context of chert
Dove P, Rimstidt J (1994) Silica-water interactions: silica:
units and their surroundings must be considered physical behavior. Geochem Mater Appl 29:259
closely. To obtain a continuous stratigraphic Fleming B, Crerar D (1982) Silicic-acid ionization and
sequence, unaltered by surface conditions and calculation of silica solubility at elevated-temperature
and ph – application to geothermal fluid processing
unaffected by lightning strikes (for paleomag-
and reinjection. Geothermics 11(1):15–29
netic studies), it will be important to drill cores Hofmann A, Harris C (2008) Silica alteration zones in the
through chert units. Such core samples should be Barberton greenstone belt: a window into subseafloor
studied collaboratively using different techniques processes 3.5–3.3 Ga ago. Chem Geol
257(3–4):224–242
and methods to obtain a more accurate insight
Kato Y, Nakamura K (2003) Origin and global tectonic
into the conditions of life in the Archean. Amor- significance of early Archean cherts from the Marble
phous silica deposits on Mars deserve full atten- Bar greenstone belt, Pilbara craton Western Australia.
tion in preparation for future missions such as Precambrian Res 125(3–4):191–243
Knauth L, Lowe D (2003) High Archean climatic temper-
ExoMars and Mars Sample Return.
ature inferred from oxygen isotope geochemistry of
cherts in the 3.5 Ga Swaziland Supergroup, South
Africa. Geol Soc Am Bull 115(5):566–580
See Also Laruelle G, Roubeix V, Sferratore A, Brodherr B,
Ciuffa D, Conley D, Durr H, Garnier J, Lancelot C,
Phuong Q, Meunier J, Meybeck M, Michalopoulos P,
▶ Alteration Moriceau B, Longphuirt S et al (2009) Anthropogenic
▶ Apex Basalt, Australia perturbations of the silicon cycle at the global scale:
▶ Apex Chert key role of the land-ocean transition. Glob
Biogeochem Cycles. 23
▶ Apex Chert, Microfossils
Maliva R, Knoll A, Simonson B (2005) Secular change in
▶ Archean Drilling Projects the Precambrian silica cycle: insights from chert
▶ Archean Environmental Conditions petrology. Geol Soc Am Bull 117(7–8):835–845
Chicken or Egg Problem 443
Nutman A, Friend C (2009) New 1:20, 000 scale geolog- microbial ecosystem: processes on the Early Earth.
ical maps, synthesis and history of investigation of the 405:105–131
Isua supracrustal belt and adjacent orthogneisses, Zegers T, de Wit M, Dann J, White S (1998) Vaalbara,
southern West Greenland a glimpse of Eoarchaean Earth’s oldest assembled continent? A combined
crust formation and orogeny. Precambrian Res structural, geochronological, and palaeomagnetic
172(3–4):189–211 test. Terra Nova 10(5):250–259
Nutman A, Allaart J, Bridgewater D, Dimroth E, Rosing
M (1984) Stratigraphic and geochemical evidence for
the depositional environment of the early Archean Isua
C
supracrustal belt Southern West Greenland. Precam- Chicken or Egg Problem
brian Res 25(4):365–396
Polat A, Frei R (2005) The origin of early Archean
banded iron formations and of continental crust, Henderson James (Jim) Cleaves II
Isua, southern West Greenland. Precambrian Res Earth–Life Science Institute (ELSI), Tokyo
138(1–2):151–175 Institute of Technology, Meguro–ku, Tokyo,
Rice M, Bell J, Cloutis E, Wang A, Ruff S, Craig M,
Japan
Bailey D, Johnson J, de Souza P, Farrand W (2010)
Silica-rich deposits and hydrated minerals at Gusev Institute for Advanced Study, Princeton, NJ,
Crater Mars: Vis-NIR spectral characterization and USA
regional mapping. Icarus 205(2):375–395 Blue Marble Space Institute of Science,
Washington, DC, USA
isotopes in cherts. Nature 443(7114):969–972 Center for Chemical Evolution, Georgia Institute
Schopf J, Kudryavtsev A, Agresti D, Wdowiak T, Czaja of Technology, Atlanta, GA, USA
A (2002) Laser-Raman imagery of Earth’s earliest
fossils. Nature 416(6876):73–76
Suganuma Y, Hamano Y, Niitsuma S, Hoashi M,
Hisamitsu T, Niitsuma N, Kodama K, Nedachi Definition
M (2006) Paleomagnetism of the Marble Bar chert
member. Western Australia: implications for apparent Chicken or egg problems are apparent causality
polar wander path for Pilbara craton during Archean
problems; for example, which came first, the
time. Earth Planet Sci Lett 252(3–4):360–371
Sugitani K, Mimuar K, Nagaokoa T, Lepot K, Takeuchi chicken or the egg? This is paradoxical because
M (2013) Microfossil assemblage from the 3400 Ma chickens are required to lay eggs, and yet chickens
Strelley pool formation in the Pilbara craton. Western hatch from eggs. An apparent paradox in modern
Australia: results form a new locality. Precambrian
biochemistry is the fact that nucleic acids (both
Res 226:59–74
Tice M, Lowe D (2004) Photosynthetic microbial mats in DNA and RNA) are required to make coded pro-
the 3, 416-Myr-old ocean. Nature 431(7008):549–552 teins, and at the same time, coded proteins are
van den Boorn S, van Bergen M, Nijman W, Vroon required to make nucleic acids. In the origin and
P (2007) Dual role of seawater and hydrothermal fluids
early evolution of life then, it is not apparent which
in early Archean chert formation: evidence from sili-
con isotopes. Geology 35(10):939–942 of these two must have come first. One possible
van den Boorn S, van Bergen M, Vroon P, de Vries S, solution to this problem is offered by the discovery
Nijman W (2010) Silicon isotope and trace element of ribozymes, RNA molecules which could poten-
constraints on the origin of similar to 3.5 Ga cherts:
tially code for their own replication and serve as
implications for early Archaean marine environments.
Geochim Cosmochim Acta 74(3):1077–1103 catalysts for their replication as well.
Van Kranendonk M, Hickman A, Smithies R, Nelson D,
Pike G (2002) Geology and tectonic evolution of the
Archean North Pilbara terrain, Pilbara Craton Western
See Also
Australia. Econ Geol Bull Soc 97(4):695–732
Westall F (2008) Morphological biosignatures in early
terrestrial and extraterrestrial materials. Space Sci ▶ Genetic Code
Rev 135(1–4):95–114 ▶ Nucleic Acids
Westall F, de Vries S, Nijman W, Rouchon V, Orberger B,
▶ Origin of Life
Pearson V, Watson J, Verchovsky A, Wright I,
Rouzaud J, Marchesini D, Severine A (2006) The ▶ Protein
3.466 Ga “Kitty’s Gap Chert,” an early Archean ▶ RNA World
444 Chicxulub Crater
central peak area. The crater was drilled first for

Chicxulub Crater oil exploration purposes. In 2002, the International
Continental Scientific Drilling Program drilled the
Philippe Claeys crater for scientific research. The cores yielded
Earth System Science, Vrije Universiteit Brussel, ▶ suevite (a breccia of melt and solid fragments
Brussels, Belgium floating in a fine pulverized clastic matrix) and
impact melt-rock containing evidence of high-
pressure metamorphism, such as ▶ shocked quartz
Keywords and diaplectic glass (i.e., formed by impact
shockwaves). The identified melt phases have a
Carbonaceous chondrites; Dinosaurs; Crater 40
Ar39Ar age of 65.46 0.50 Ma, corresponding
(impact); Mass extinctions; Meteorites to the age of the KT boundary. The ▶ impactites
are linked geochemically to the ▶ ejecta material
spread all over the Gulf of Mexico. The energy
Definition liberated by the impact amounted to 1,024 J. The
crater excavation injected into the upper atmo-
The Chicxulub structure is an impact crater sphere vast quantities of water vapor (200 Gt),
formed 65 Ma ago and located at the tip of the CO2 (350–3,500 Gt), and SOx (40–560 Gt) pro-
Yucatan Peninsula (Mexico). With a diameter duced by the vaporization of the target material,
between 180 and 220 km, the Chicxulub structure along with a huge volume of fine dust. The dust
is the third largest impact crater known on Earth. and sulfur aerosols led to a yearlong darkness and
The Chicxulub impact is considered by many brutal cooling all over the Earth, strongly reducing
authors as a cause of the important biological photosynthesis. Climate models suggested that a
crisis at the ▶ KT boundary that led to mass CO2-induced greenhouse warming followed.
extinction of 50–60 % of the fauna and flora on These events likely caused or contribute to the
the continent and in the oceans, including organ- important biological crisis precisely at the ▶ KT
isms such as ammonites and in particular the boundary that led to the mass extinction of more
non-avian dinosaurs. than 60 % of the fauna and in particular of the
non-avian dinosaurs.
Overview
See Also
The Chicxulub structure formed 65 million years
▶ Crater, Impact
ago by the collision of a projectile, most likely a
▶ Deccan Trapps
10–12 km in size ▶ carbonaceous chondrite mete-
▶ Ejecta
orite, with the Earth crust. The impact took place
▶ Impactite
on a shallow-water carbonate platform; the under-
▶ KT Boundary
lying target rocks were composed of 2–3 km of
▶ Mass Extinctions
sediments (carbonates and ▶ evaporites) overly-
▶ Shocked Quartz
ing a 600-Ma-old gneiss-granitic basement. Geo-
▶ Suevite
physical methods give a good image of the crater
that lies buried under 1 km of Cenozoic sedi-
ments. Seismic data show the concave morphol- References and Further Reading
ogy of the structure. Measurements of the gravity
field clearly outline a central-peak ring surrounded Morgan J, Warner M (1999) Chicxulub: the third dimen-
sion of a multi-ring impact basin. Geology 27:407–410
by a region of lower density forming the trough Pierazzo E et al (2003) Chicxulub and climate: radiative
zone. A magnetic anomaly, caused by the presence perturbations of impact-produced S-bearing gases.
of iron-rich rocks, corresponds to the uplifted Astrobiology 3:99–118
Chirality 445
Schulte P et al (2010) The Chicxulub asteroid impact and element of symmetry (excepting in some cases an
mass extinction at the cretaceous-paleogene boundary. axis of rotation). The two distinct mirror-image
Science 327:1214–1218
Swisher CC et al (1992) Coeval 40Ar/39Ar ages of 65.0 forms are called enantiomers in the case of chiral
million years ago from Chicxulub crater melt rock and molecules and enantiomorphs in the case of chi-
Cretaceous-Tertiary boundary tektites. Science ral crystals.
257:954–958
Urrutia-Fucugauchi J et al (2004) The Chicxulub scientific
drilling project (CSDP). Meteorit Planet Sci
C
39:787–790 Overview
The two enantiomers of a chiral molecule have

identical physical and chemical properties, which
Chiral Excess was demonstrated by van’t Hoff and Le Bel using
arguments about the “exact mechanical symme-
▶ Enantiomeric Excess try” between enantiomers (van’t Hoff 1887). It
has since been proven, however, that because of
parity violation, there is a minute – and, to date,
experimentally immeasurable – energy differ-
Chirality ence between enantiomeric molecules, which by
current predictions lie in the range of femto- to
Donna Blackmond picojoules/mol (Quack 2002).
The Scripps Research Institute, La Jolla, CA, USA Each of the two enantiomeric forms of a mol-
ecule interacts differently with other chiral mol-
ecules, chiral objects, or chiral forces such as
Keywords plane-polarized light. Such chiral interactions
form the basis for the molecular recognition that
Enantiomers; Enantiomorphs; Diastereomers; enables the specificity of reactions observed in
Stereochemistry; Conglomerates; Racemic com- biological systems.
pounds; Symmetry breaking; Chiral bias; Chiral A collection of enantiomers of a molecule may
amplification; Biological homochirality be comprised of equal amounts of each hand
(a racemic mixture), all of a single hand
(an enantiopure mixture), or any fraction of each
Synonyms enantiomer (a scalemic mixture). The standard
definition of enantiomeric purity, known as the
Handedness; Non-superimposable mirror image enantiomeric excess, arose from the observation
object that pure enantiomers rotate the plane of polarized
light to the same degree but in opposite directions.
Thus, an equal mixture of left- and right-handed
Definition molecules exhibits no rotation of plane-polarized
light. The enantiomeric excess of a collection of
The term chirality, from the Greek word cheir for enantiomeric molecules is defined by the degree
“hand,” refers to the property of “handedness” and direction of rotation compared to the
possessed by some molecules and macroscopic enantiopure case and is given by Eq. 1:
objects. A molecule or object is chiral if it is not
superimposable on its mirror image, as is the case RS
%enantiomeric excess ¼ %ee ¼ 100
for left and right hands. The word was introduced RþS
by Lord Kelvin (1904) and has been further quan-
tified by Mislow (1965) and Cahn et al. (1966), where R and S are defined in the following
who define an object as chiral when it contains no paragraph.
446 Chirality
The term homochirality typically refers to the single chirality is a central problem that has
fact that the two main building blocks of biolog- intrigued scientists for generations. It is in fact
ical molecules, sugars and amino acids, each two questions (Blackmond 2009): First, how was
exist in only one of the two possible enantiomeric an initial chiral bias induced, known as symmetry
forms. Convention describes these as right- breaking? And second, how was such an imbal-
handed, R, or d-sugars and left-handed, S, or ance sustained and propagated to high levels of
l-amino acids. The term homochirality was intro- enantiopurity?
duced by Lord Kelvin, who wrote: “two equal Proposals for how an imbalance initially came
and similar right hands are homochirally similar” about may be classified as either terrestrial or
(Lord Kelvin, 1904). This description could extraterrestrial and then subdivided into either
apply, for example, to a series of different random or deterministic, sometimes called “de
amino acids with all in the same enantiopure facto” and “de lege,” respectively (Quack
form or to an enantiopure collection of molecules 2002). Evidence of small enantiomeric excesses
of a single amino acid. Biopolymers formed from in amino acids found in chondritic meteor
d-sugars, such as RNA and DNA, and from deposits has been suggested to support the extra-
l-amino acids, such as polypeptides and proteins, terrestrial breaking of symmetry (Burton
are homochiral. et al. 2012). The possible role of parity violation
In the modern literature concerning origin of energy differences fits into a “de lege” model,
life, the term homochirality has become synony- although to date no chemical or physical differ-
mous with the single-handedness of the molecu- ence between enantiomers has been confirmed in
lar building blocks and biopolymers of living experiments.
systems. Homochirality has been suggested as a Proponents on the “chance” side of this ques-
possible biosignature, an idea that has been tion suggest that symmetry breaking could occur
expressed even more strongly as “the homochiral stochastically due to transient fluctuations in the
imperative of molecular evolution” (Siegel 1998) physical and chemical environment. However,
or by the tenet of “No homochirality, no life!” any small imbalance created in this way should
(Bonner 1998). More recent experiments demon- average out as the racemic state unless some
strating that a d-polymerase ribozyme could process intervenes to sustain and amplify
evolve to catalyze the cross-chiral replication of it. Thus whether or not the imbalance in enantio-
its mirror image suggest the possibility that rep- mers came about by chance, arising on Earth or
licating d- and l-RNA molecules may have elsewhere, an amplification mechanism remains
emerged together Sczepanski and Joyce 2014). the key to increasing enantiomeric excess.
In such a scenario, the homochiral path of modern Theoretical work from the 1950s (Frank 1953)
life might have arisen from later evolutionary and experimental studies from the 1990s to the
pressures on early forms of life. present day have provided a number of proposals
for the amplification of a small initial imbalance in
Emergence of Homochirality enantiomers. Experimental models invoke both
The abiotic synthesis of chiral molecules results chemical reactions and physical phase behavior
in a racemic mixture of products in the absence of to rationalize enantioenrichment. Chemical
a chiral bias that helps to direct the reaction models invoke autocatalytic reactions, chirality
preferentially toward one hand. In modern transfer, or kinetic resolutions, while physical
organic synthesis, such a chiral bias may be models exploit aspects of the crystallization
obtained from the pool of enantioenriched mole- behavior of chiral molecules to provide enantioen-
cules produced by living systems. In a prebiotic richment of either the solid or solution phases.
world, however, prior to the presence of a chiral
pool, racemic mixtures of chiral molecules would Chemical Models for Enantioenrichment
be expected from such reactions. How a racemic The Soai reaction provides the most robust
prebiotic world came to develop life based on experimental example of asymmetric
Chirality 447
autocatalytic amplification of enantiomeric The majority of chiral molecules – including

excess (Soai et al. 1995). Although the chemis- 17 of the 19 proteinogenic amino acids that are
try is not itself prebiotically plausible, the reac- chiral – do not crystallize as conglomerates or
tion provides the first experimental proof of separate left- and right-handed crystals, but
concept of the Frank model for the emergence instead form heterochiral crystals usually com-
of homochirality (Blackmond 2004). Catalysis prised of a 1:1 ratio of enantiomers, called race-
of the formose reaction by amino acids (Breslow mic compounds. Models for enantioenrichment C
and Cheng 2010) or amino acid salts (Hein and based on the phase behavior of racemic com-
Blackmond 2012) leads to enantioenrichment of pounds rely on the thermodynamic partitioning
the resulting glyceraldehyde, the smallest chiral of the enantiomers between the solution and solid
sugar. The transfer of chirality to amino acids phases. A system containing an unequal number
via transamination reactions has been demon- of left-and right-handed molecules in an equilib-
strated (Levine et al. 2008). A kinetic resolution rium mixture of crystals in saturated solution
of sugars by amino acids – or, conversely, of contains both the heterochiral solid and an
amino acids by sugars – results in significant enantiopure solid of the enantiomer present in
enantioenrichment (Hein et al. 2011) of precur- excess. The composition of the solution phase –
sors in a prebiotically plausible route to RNA that is, the solution enantiomeric excess – is a
(Powner et al. 2009). constant value dictated by the Gibbs phase rule.
Known as the eutectic composition, this is a
Physical Models for Enantioenrichment characteristic property of each amino acid and is
Ever since Pasteur used tweezers to separate the related to the relative solubilities of the
mirror-image crystals of the di-salt of tartaric heterochiral and homochiral solids. For example,
acid, crystallization processes and the phase serine and valine exhibit eutectic compositions of
behavior of chiral molecules have intrigued sci- 99 % ee and 47 % ee, respectively. This property
entists as a potential means for the emergence of was first recognized as a possible route to
homochirality. Kondepudi showed striking enantioenrichment by Morowitz (1969) and was
amplification of the enantiomeric excess of elaborated by Blackmond and coworkers
NaClO3 crystals by rapidly stirring supersatu- (Klussmann et al. 2006a, b) and by Breslow and
rated solutions (Kondepudi 1990), and Viedma coworkers (Breslow and Cheng 2009). A similar
showed that racemic mixtures of these same crys- approach to enantioenrichment via sublimation
tals evolved to homochirality under the influence of amino acids has been reported (Fletcher
of attrition (Viedma 2005). The former case is et al. 2007; Perry et al. 2007).
noted as a “far from equilibrium” process, while The explosion of experimental reports on the
the latter takes place close to equilibrium; subject of the emergence of biological
both rely on the fact that while NaClO3 is homochirality in the first decade of the twenty-
achiral in solution, it forms separate left- and first century led Chemistry World to suggest that
right-handed crystals. It was recognized that we are now “spoilt for choice” (Ball 2007) in
such processes for chiral amplification might be proposals for answering the puzzle of the single
extended to intrinsically chiral molecules that chirality of life on Earth.
form separate crystals (called conglomerates) if
the enantiomers could be made to exchange rap-
idly in solution. The first proof of concept of See Also
attrition-enhanced deracemization of intrinsi-
cally chiral molecules was demonstrated for ▶ Enantiomeric Excess
amino acid derivatives (Noorduin et al. 2008) ▶ Enantiomers
and for a proteinogenic amino acid ▶ Homochirality
(proteinogenic refers to amino acids that are pre- ▶ Origin of Life
cursors to proteins) (Viedma et al. 2008). ▶ Racemic Mixture
448 Chiron
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Lett 10:2433–2436 an ▶ asteroid, labeled 2060 Chiron. In 1989, as the
Lord Kelvin (W. Thomson) (1904) Baltimore lectures on object was moving toward perihelion, a coma was
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discovered and it was also designated as a comet,
C. J. Clay and Sons, London, p 619 (The lectures
were presented in 1884 and 1893 at Johns Hopkins 95 P/Chiron. Chiron is the first identified object
University, Baltimore) among the class of the ▶ Centaurs which orbit
Chlorophylls 449
between Jupiter and Neptune. Its period is 50 years See Also

and its perihelion distance is 8.4 AU. Its diameter
is about 180 km and its albedo is about 0.05. ▶ Diffuse Cloud
Chiron is now considered to belong to the C-type ▶ Herschel Mission
class of asteroids. Like other Centaurs, Chiron is ▶ Hydrogen Chloride
assumed to be originally a ▶ trans-Neptunian ▶ Interstellar Medium
object which escaped from the ▶ Kuiper Belt. ▶ Isotopolog C
See Also
▶ Kuiper Belt Chloronium
▶ Trans-Neptunian Object
▶ Chlorine Hydrides in the Interstellar Medium
Chlorine Hydrides in the Interstellar

Medium
Chloroniumyl
William M. Irvine
University of Massachusetts, Amherst, MA, USA ▶ Chlorine Hydrides in the Interstellar Medium
Synonyms
Chloronium; Chloroniumyl; H2Cl+; HCl; HCl+; Chlorophylls

Hydrogen chloride
Juli Peretó
Definition Spain
Not only the familiar diatomic molecule

containing hydrogen and chlorine, ▶ hydrogen Definition
chloride (HCl), but also the cations HCl+ and
H2Cl+ have been detected by astronomers in the Chlorophylls are a class of pigments derivative of
▶ interstellar medium. In fact, both the 35Cl and protoporphyrin complexed with magnesium.
the 37Cl ▶ isotopologs for all three molecular Chlorophylls function in photosynthetic organ-
species have been identified in space. isms both as light receptors and special photo-
chemical devices in photosynthetic reaction
centers.
History
Whereas HCl was detected using the Kuiper Air- See Also
borne Observatory (KAO), HCl+ and H2Cl+ were
observed with the Herschel Space Observatory. ▶ Bacteriochlorophyll
For more information, see ▶ Molecules in Space. ▶ Photosynthesis
450 Chloroplast
autonomy, size, and morphological resemblance

Chloroplast to prokaryotes, it was suggested that chloroplasts
were descendants of bacteria by ▶ endosymbiosis.
Ricardo Amils There is different structural, functional, and molec-
Departamento de Biologia Molecular, ular evidence that supports the endosymbiotic ori-
Universidad Autónoma de Madrid, Madrid, gin of chloroplasts: existence of a small genome,
Spain presence of ribosomes, antibiotic specificity, and
molecular phylogeny.
Definition
See Also
Chloroplast is a ▶ chlorophyll-containing organ-
elle of phototrophic eukaryotes responsible for the ▶ Algae
generation of energy from radiation. A major fea- ▶ Anoxygenic Photosynthesis
ture of eukaryotic cells, absent from prokaryotic ▶ ATP Synthase
cells, is the presence of membrane-enclosed struc- ▶ Autotrophy
tures called ▶ organelles. These include mitochon- ▶ Calvin-Benson Cycle
dria and chloroplasts, the latter only in ▶ Carbon Dioxide
photosynthetic cells. Like mitochondria, chloro- ▶ Chlorophylls
plasts have a permeable outermost membrane, a ▶ Endosymbiosis
much less permeable inner membrane, and an ▶ Organelle
intermembrane space. The inner membrane sur- ▶ Photoautotroph
rounds the lumen of the chloroplast, called stroma. ▶ Photosynthesis
Chlorophyll and all other components needed for ▶ Photosynthetic Pigments
▶ photosynthesis are located in a series of flattened ▶ Proton Motive Force
membrane disks called thylakoids. The thylakoid
membrane is highly impermeable to ions and other
metabolites because its function is to establish the
▶ proton motive force necessary for ATP synthe- Chondrite
sis. In green ▶ algae and plants, thylakoids are
typically stacked into discrete structural units Frank Sohl
called grana. The chloroplast stroma contains Deutsches Zentrum f€ur Luft- und Raumfahrt
large amounts of the enzyme ribulose bisphosphate (DLR), Institut f€ur Planetenforschung, Berlin,
carboxylase (RubisCO). RubisCO is a key catalyst Germany
of the Calvin cycle, the series of biosynthetic reac-
tions by which most photosynthetic organisms
convert CO2 to organic compounds. RubisCO Definition
makes up over 50 % of the total chloroplast protein
and catalyzes the formation of phosphoglyceric Chondrites are undifferentiated stony ▶ meteor-
acid, a key compound in the biosynthesis of glu- ites and represent the majority among stony
cose. The permeability of the outermost chloro- meteorites. The term literally means “with
plast membrane allows glucose and ATP ▶ chondrules” and therefore underlines the main
produced during photosynthesis to diffuse into difference to ▶ achondrites. Chondrites consist of
the cytoplasm where they can be used to build a fine-grained matrix of micrometer-sized dust
new cell material. On the basis of their relative particles, surrounding roughly millimeter-sized,
Chromatographic Coelution 451
round inclusions (▶ Chondrule), refractory roughly millimeter-sized silicates. Their forma-

▶ CAIs (Ca-Al-rich inclusions), particles enriched tion is not fully understood, but they did go
in metallic Fe-Ni and sulfides, and other individual through a transient heating process which was
mineral grains. The matrix also contains presolar followed by a cooling period, so that molten or
grains that formed elsewhere in the Galaxy before partly molten droplets came together before these
the Solar System came into being. The presence of pieces accreted to their parent body. Chondrules
▶ chondrules may suggest that chondrites were together with the ▶ CAIs (Ca-Al-rich inclusions) C
formed by accretion of thermally processed dust constitute the oldest material of our
particles, whereas their primitive parent bodies ▶ Solar System.
never experienced partial melting and recrystalli-
zation. The class of chondrites has been divided
into enstatite (EH and EL types), Rumuruti See Also
(R type), ordinary (H, L, and LL types), and
carbonaceous (C types); the latter is subject to ▶ CAIs
aqueous alteration and thought to be the most ▶ Chondrite
primitive meteorites. H, L, and LL indicate high ▶ Meteorites
iron, low iron, and low iron/metal contents, ▶ Silicate
respectively. ▶ System Solar Formation, Chronology of
See Also
▶ Achondrite Chromatographic Coelution

▶ CAIs
▶ Chondrule Jason P. Dworkin
▶ Meteorites NASA Goddard Space Flight Center,
Astrochemistry Laboratory, Greenbelt, MD,
USA
Chondrule Synonyms
Frank Sohl and Tilman Spohn Co-elution

(DLR), Institut f€
ur Planetenforschung, Berlin,
Germany Definition
Chromatographic coelution occurs when two

Definition (or more) compounds do not chromatographi-
cally separate due to the fact that both species
The term chondrule comes from the Greek word have retention times that differ by less than the
“chondros” for grain or seed. They are the major resolution of the method. This can be solved by
component of ▶ chondrites and can make up to increasing the selectivity and/or efficiency of the
80 % of the volume of a given ▶ meteorite. In ▶ chromatography or by employing different
general, chondrules are rounded inclusions, detection techniques that can differentiate the
452 Chromatography
L-Aspartic Acid D Glutamic Acid

D-Aspartic Acid L Glutamic Acid
Chromatography
Jason P. Dworkin
B
NASA Goddard Space Flight Center,
D+L Aspartic Acid
Astrochemistry Laboratory, Greenbelt, MD, USA
Intensity
D+L Glutamic Acid
Keywords
Column chromatography; GC; Hyphenated tech-

niques; LC; Planar chromatography; Separation
A science; Solid phase extraction
Time
Definition
Chromatographic Coelution, Fig. 1 Chromatographic
co-elution can be resolved by adjusting the chromato-
Chromatography (either preparative or analyti-
graphic conditions. In these reverse phase HPLC traces,
fluorescently labeled amino acids are resolved by decreas- cal) is a qualitative or quantitative experimental
ing the methanol concentration in the mobile phase from method based on the properties of all molecules
trace A to trace B to partition more or less selectively from one
phase into another and therefore to migrate at
different rates when they are carried across a
coeluting compounds. Changing the chemistry of
solid or liquid stationary phase by a mobile
the mobile phase, stationary phase, temperature,
phase, which can be a gas, liquid, or supercritical
and column or plane length are good methods to
fluid.
increase the separation. Techniques such as mass
spectrometry and optical spectroscopy are com-
mon ways to distinguish between coeluting com-
Overview
pounds that cannot be resolved (Fig. 1).
Chromatography is a broad laboratory method for
physically separating compounds from a mixture
See Also
based on the preferential partitioning of different
molecules across two different phases. The ana-
▶ Chromatography
lyte is carried in the mobile phase (gas, liquid, or
▶ Gas Chromatography
supercritical fluid) and then passes through a sta-
▶ GC/MS
tionary phase (liquid or solid), and some analytes
▶ Ion-Exchange Chromatography
are retained more efficiently than others. Differ-
▶ Liquid Chromatography-Mass Spectrometry
ent types of chromatography are based on the
▶ Pyrolysis GC/MS
geometry (through a stationary phase-filled tube
or “column” or over a plate or “plane” with sta-
References and Further Reading tionary phase), the type of mobile/stationary
phase interaction (gas/liquid, gas/solid, liquid/
Burton AS, Stern JC, Elsila JE, Glavin DP, Dworkin JP liquid, liquid/solid, supercritical fluid/solid), and
(2012) Understanding prebiotic chemistry through the
analysis of extraterrestrial amino acids and
the nature of the analyte/stationary phase interac-
nucleobases in meteorites. Chem Soc Rev tion (e.g., host/guest chemistry, hydrophobic
41:5459–5472 interactions, ion exchange, hydrodynamic
Chromophore 453
volume, volatility, etc.). ▶ Electrophoresis, while ▶ Gas Chromatography

not strictly chromatography, is closely related. ▶ GC/MS
While planar chromatography (e.g., paper and ▶ Ion-Exchange Chromatography
thin layer (TLC)) is typically qualitative, column ▶ Liquid Chromatography-Mass Spectrometry
chromatography is well suited for real-time inter- ▶ Mars Science Laboratory
rogation of the eluting compounds with detectors. ▶ Mass Spectrometry
Common types of gas-liquid chromatography ▶ Pyrolysis GC/MS C
(GC) detection techniques are flame ionization ▶ Viking
detection, thermal conductivity, and any number
of mass spectrometric techniques (GC-MS).
Common detectors in liquid/solid chromatogra- References and Further Reading
phy (LC or ▶ HPLC) are ultraviolet absorbance,
fluorescence, refractive index, and various mass Ettre LS (1993) Nomenclature for chromatography. Pure
Appl Chem 65:819–872, http://chromatogra-
spectrometric methods (LC-MS). The addition of
phyonline.com/
more elaborate detectors with a chromatographic Miller JM (2004) Chromatography: concepts and con-
or capillary electrophoretic “front end” trasts. Wiley-Interscience, Hoboken, p 490
(or “inlet”) is collectively referred to as hyphen-
ated or hybrid techniques. Combinations of chro-
matography with ▶ mass spectrometry, nuclear
magnetic resonance, and optical spectroscopy
increase the ability of an analytical chemist to Chromophore
study small quantities of complex mixtures.
Conversely, preparative chromatography John H. Chalmers
(either column or planar) is used to purify large Scripps Institute of Oceanography Geosciences
quantities of a compound of interest. While GC or Research Division, University of California, San
supercritical fluid chromatography is sometimes Diego, La Jolla, CA, USA
used, the most popular method is LC
(or electrophoresis). One popular method is
solid-phase extraction (SPE). Definition
While the majority of chromatography is
conducted in laboratories or industry, the process Chromophores are chemical groups that absorb
can be found elsewhere. Naturally occurring ▶ electromagnetic radiation, for example, in the
chromatography is typically fractional distilla- visible, near-infrared, or ultraviolet ranges, though
tion or geochromatography, where, for example, the original derivation of the word comes from the
compounds dissolved in water are roughly sepa- Greek for color, implying absorption of visible
rated as they pass through rocks or sediment radiation. They may contain systems of conju-
columns. Chromatography (GC-MS) has also gated multiple bonds, transition element ions, or
been used in robotic planetary science missions both. In biological systems, light absorption by
to study Mars; Saturn’s moon, Titan; and comets. chromophores forms the basis of vision, ▶ photo-
synthesis, and, in the case of ultraviolet light,
▶ mutation and vitamin D synthesis. In the context
of astrobiology, chromophores have been
See Also found, for example, in simulated cometary ices.
Chromophores are responsible for the colors of
▶ Affinity Chromatography dyes, pigments, minerals, and many organisms,
▶ Electrophoresis though some colors, particularly blues and irides-
▶ ExoMars cent effects, are produced by diffraction.
454 Chromosome
See Also See Also
▶ Bacteriochlorophyll ▶ DNA
▶ Chlorophylls ▶ Genome
▶ Circular Dichroism ▶ Nucleus
▶ Electromagnetic Radiation ▶ Replication (Genetics)
▶ Extreme Ultraviolet Light ▶ Transcription
▶ Fluorophore
▶ Mutation
▶ Photosynthesis
▶ Photosynthetic Pigments
Chronostratigraphy
Stephan van Gasselt and Gerhard Neukum

Planetary Sciences and Remote Sensing, Institute
of Geological Sciences, Freie Universität Berlin,
Chromosome
Berlin, Germany
Juli Peretó
Institut Cavanilles de Biodiversitat i
Keywords
Biologia Evolutiva, Universitat de València,
València, Spain
Cratering chronologies; Impact crater; Produc-
tion function; Radiometric ages; Stratigraphy
Definition Synonyms
Chromosome is a structure consisting of or Cratering chronology; Planetary chronostra-

containing ▶ DNA with essential genetic infor- tigraphy; Planetary surface ages
mation for the cell. In most prokaryotes, it is a
circular, double-stranded DNA, normally
attached to the cell membrane, with a folded Definition
structure (also known as nucleoid) with some
attached proteins. In eukaryotic cells, usually The term chronology with respect to planetary
the number of chromosomes is characteristic surfaces generally refers to the timing of events
of the species – polyploid plants are a remark- when rock surface units were formed or modified
able exception. Somatic cells possess two sets and describes the position of a geologic unit
of homologous chromosomes (diploid number within its stratigraphic context. The term chro-
or 2n), one originating from the male progeni- nology furthermore encompasses the method of
tor, the other from the female. In contrast, germ- deriving ages of planetary surface units by means
line cells are haploid (n). Eukaryotic chromo- of radiometric age determinations as well as by
somes are located inside the ▶ nucleus and con- analyses of impact crater size-frequency distribu-
sist of a double-stranded DNA molecule highly tions based on knowledge or assumptions of the
folded and complexed with proteins (histones) flux and size distribution of planet-impacting
also known as chromatin. The term chromo- bodies in the history of the Solar System.
some also applies to the DNA molecules located
in mitochondria and plastids and to the genome
of DNA viruses.
Gerhard Neukum: deceased.
Chronostratigraphy 455
Earth Moon Mercury Venus Mars
.5 .5
1 Ga 1 Ga
C
.5 .5
2 Ga 2 Ga
.5 .5
3 Ga 3 Ga
.5 .5
4 Ga 4 Ga
.5 .5
Chronostratigraphy, Fig. 1 Chronology units in terms 5, Neukum et al. (2001a), 6, Strom and Neukum (1988),
of epochs and periods for the terrestrial planets as com- 7, Spudis and Guest (1988), 8, Basilevsky and Head
piled from different sources; see main text for description; (1998), 9, Basilevsky and Head (2002), 10, Tanaka
the two columns for Venus’ chronology depict the upper (1986), 11, Neukum and Wise (1976), 12a, Hartmann
and lower estimates for T = 800 Ma and T = 288 Ma, and Neukum (2001) Neukum model, and 12b, Hartmann
respectively (1, Ogg et al. (2008), 2, Neukum and Ivanov and Neukum (2001) Hartmann model)
(1994), 3, Stöffler and Ryder (2001), 4, Wilhelms (1987),
Overview refers to an age derived by various methods as

discussed below, a chronostratigraphic unit con-
The chronology of planetary surfaces refers to the sists of rocks formed on a global scale during a
sequence of geological events that have led to specific geologic time (Ogg et al. 2008).
deposition and emplacement of rock-forming The knowledge we have today about the
units on a planetary surface. The sequence of chronostratigraphy and chronology of terrestrial
events and the geologic materials, i.e., the rock- planetary surfaces other than the Earth is based
forming units (Wilhelms 1987; Greeley and upon the knowledge we have gained during plan-
Batson 1990), are the planet’s chronostra- etary exploration of the ▶ Moon (Wilhelms
tigraphic record and have to be seen in a specific et al. 1971) and was later expanded to other
planetary context as each planetary body evolves terrestrial surfaces, in particular ▶ Mercury and
in a different way. ▶ Mars for which formal stratigraphic systems
While the chronology in general refers to the have been developed (Tanaka and Hartmann
timing and sequence of such events, the strati- 2008).
graphic record of a planet is defined by divisions The establishment of timescales for planetary
of geologic time and boundaries of geologic units surfaces is based upon geological mapping of
on a global scale. Divisions of time are charac- planetary surfaces as defined by surface proper-
terized by primary or secondary markers consid- ties, e.g., morphologies, textures, spectral com-
ered to be characteristic of that surface. While the positions, and, as delineated by geologic
geologic time unit (chronologic unit; see Fig. 1) contacts, the superposition of individual units
456 Chronostratigraphy
following basic stratigraphic principles. Relative units and their relation to neighboring units.
ages can be derived by measuring the size- Ages of rocky planetary surface material are
frequency distribution of ▶ impact craters that determined by means of the decay of radioactive
have accumulated in a planet’s history through isotopes and decay products if decay rates are
time, which means that densely cratered surfaces known and if samples of surface material are
are older than less-densely cratered surface units. available. In cases where planetary surfaces are
Thus, a geologic unit records the age as expressed observed by remote-sensing methods, ages of
by the number of impact craters formed during surface forming units can only be determined by
meteoritic bombardment and the time a unit was analyses of impact crater size-frequency mea-
exposed to the projectile impact flux (Öpik 1960; surements, i.e., by comparing the size-frequency
Baldwin 1964; Hartmann 1966; Neukum distribution of impact craters with the modeled
et al. 1975). size distribution and impact flux of meteorites on
Absolute ages for surface units are constrained a given planetary surface.
by samples returned from the Moon in the course While relative age determinations of units
of the lunar Apollo and Luna mission programs. allow to assess and estimate the relative sequence
Derived radiometric ages of surface samples have of unit-forming events within an area of interest
established a calibration for the size-frequency and in comparison with an established global
distribution of the Moon’s surface as observed distribution of impact crater size-frequency
by remote-sensing imaging. This process allows observations, age determinations of geologic
us to measure model ages for other planetary units based on remotely sensed data with the
surfaces either (a) by estimating the relative help of either appropriate scaling laws or model-
cratering rates in comparison with the Moon or ing of impactor flux allow us to derive absolute
(b) by directly assessing cratering rates under ages. Except for the Earth, only the Moon has
consideration of impact probabilities and impac- been directly sampled thus far and therefore
tor sizes and sources (Neukum and Ivanov 1994; forms the only natural calibration target for
Tanaka and Hartmann 2008) as well as crater establishing planetary chronostratigraphies and
scaling laws that describe the relationship surface chronologies.
between projectile size and impact condition
parameters, e.g., velocity, angle, density, and sur- Radiogenic Isotope Measurements
face parameters, such as surface gravity, density, Radioactive (or radiometric) dating, i.e., determi-
compositions, and strength of surface materials nation of radiogenic isotope ages of rock samples
(Croft 1985; Holsapple 1987; Schmidt and in the context of chronology, makes use of the
Housen 1987). constancy of rates of radioactive decay by which
For planetary surface chronologies, the num- a radioactive nuclide is transformed to its daugh-
ber of stratigraphic systems for planetary objects ter product. Radiogenic isotope ages for lunar
has advanced with new data that have become material have been derived using decay measure-
available in the context of planetary exploration ments of Rb-Sr, Sm-Nd, and Ar40-Ar39. Such
mission. Stratigraphic systems cover the terres- methods are described in detail by, e.g., Dalrym-
trial (inner) planets as well as the Moon and also ple and Ryder (1991) and Albarède (2009).
the major icy satellites of the Jovian and Satur- For the Moon, over 380 kg of rock material in
nian planetary systems. over 2,000 samples was returned to Earth during
the six manned Apollo missions. During the
Soviet robotic Luna missions, several hundred
Basic Methodology grams were collected during three Luna missions
and analyzed after return to Earth. Samples are
There are different methods of assessing the stra- mainly from basaltic rock and impact glass from
tigraphy of planetary surfaces with respect to the the lunar ▶ mare areas (Apollo 11, 12, 15, 17;
relative or absolute age of individual geologic Luna 16, 24) as well as from highland terrain
(Apollo 14–17, Luna 20). In addition, a number frequency measured on a specific geologic unit is
of meteorites found on Earth have a lunar origin representative of the relative age or crater reten-
(or Martian) and could be radiometrically dated tion age of that unit (Arvidson et al. 1979).
in laboratory measurements; their exact location A geologic unit or rock-stratigraphic unit is a
of origin is, however, unknown. morphologically distinct entity formed at a spe-
Lunar rocks on the surface are exposed to cific time by a distinct geologic process (Stöffler
cosmic weathering processes and recurrent mete- et al. 2006). The derived relative surface age does C
oritic bombardment and therefore are altered. not necessarily reflect the true relative age when a
This leads to reprocessing of rocky material and geologic unit was formed, as subsequent pro-
to formation of second- or third-generation rocks cesses, generally termed resurfacing, might have
(Stöffler et al. 2006). By making use of radioiso- led to eliminating impact craters in a given
tope measurements, different ages are usually diameter-size range. The main problems arise
derived, i.e., (a) crystallization ages, (b) ages of from accurately identifying and delineating a geo-
formation of impact breccia, and (c) exposure logic unit at a given scale, as this significantly
ages. The crystallization age gives the age of depends on the appearance of a particular unit
events that led to formation of rock minerals by and its relation to surrounding units in the image
magmatic or in situ melting processes. Ages for data used for photogeologic interpretations
formation of impact breccia give insight into the (Shoemaker and Hackman 1962; Wilhelms 1987).
age rock material and minerals were transformed By employing cumulative crater size-
to breccias, while the exposure age of rock mate- frequency diagrams, the measured impact crater
rial gives the age since the rock was exposed to diameters of all impact craters within a given
cosmic weathering by cosmic rays (Stöffler geologic unit are usually plotted against their
et al. 2006). It has, however, been shown that cumulative frequency in a log-log diagram. The
direct radiometric age determinations are only impact crater size-frequency distribution derived
possible for lunar mare basalts, as highland mate- in such a way provides a measure for the surface
rial collected at the lunar surface cannot be age with older units, i.e., units with increasing
related directly to its source area. None of the crater retention ages, shifted upwards on the fre-
collected samples were derived from bedrock quency axis (apparent shift toward larger impact
material due to a several meters thick surface crater diameters), while distributions of younger
coverage of lunar regolith (e.g., Heiken units are shifted downwards on the frequency
et al. 1991 and references therein). axis leading to an apparent shift toward smaller
Radiogenic isotope measurements for deriv- impact crater diameter sizes (Hartmann
ing absolute ages of localized samples help to et al. 1981; Wilhelms 1987).
establish boundary age values, but without a care- While relative surface age determinations are
ful interpretation of the photogeological settings based upon the interpretation of remotely sensed
and the geological context, such measurements data by means of determining the superposition
cannot contribute to establishing a planet-wide of individual rock surface units and by determin-
chronology and chronostratigraphic system. ing impact crater size frequencies, age determi-
nations leading to absolute age values need
Relative Age Determinations additional pieces of information.
The easiest accessible approach to assess the time
sequence of rock-forming events on a planetary Absolute Age Determinations
surface is to measure the size-frequency distribu- There are two methods of obtaining absolute ages
tion of impact diameters within a geologic unit and of rock surface units of planetary bodies. One
to compare this to other geologic units of the same method links radiogenic isotope ages obtained
planetary body: the higher the number of impact for the Moon with crater retention ages, i.e., rel-
craters that have accumulated over time per area, ative ages, obtained through photogeological
the older the surface, which means that a crater mapping (Hartmann et al. 1981; Neukum 1983;
Neukum and Ivanov 1994). In the other approach, characterized by a terminal lunar cataclysm rather
models for rates for formation of impact craters than by a smooth, exponential decay in impact rate
are employed and are transferred to other plane- with time. The lunar cataclysm theory has been
tary objects considering their specific environ- challenged by dynamic, geologic, and strati-
ment, e.g., position and size, atmosphere, and graphic arguments (Wetherill 1975; Neukum and
target properties (Neukum and Wise 1976; Ivanov 1994; Baldwin 2006).
Neukum and Hiller 1981; Hartmann et al. 1981; Absolute ages obtained with an impact chro-
Croft 1985; Holsapple 1987; Schmidt and nology model are generally termed cratering
Housen 1987). model ages, and their units are given in gigayears
For the ▶ terrestrial planets in the Inner Solar (Ga = 1 billion years) or megayears
System, surface ages can only be obtained by (Ma = 1 million years).
models of the crater forming rates on each one Crater size-frequency distributions obtained
of these bodies. Shapes of crater size-frequency from crater size-frequency measurements are
distributions (SFDs) measured on the terrestrial represented using several techniques, of which
planets, including the Moon, were shown to be three are commonly adopted. In principle, crater
more or less similar which indicates (a) the same size diameters are grouped into pseudo-
family of bodies, preferentially asteroids (Main logarithmic or logarithmic bins and plotted on
Belt, Near-Earth asteroids, etc.), impacting these the abscissa (Arvidson et al. 1979; Neukum
planets, and (b) that time dependences of impact et al. 2001a; Michael and Neukum 2010). The
and cratering rates are similar to that for the ordinate gives the crater size-frequency values as
Moon (Neukum and Hiller 1981; Neukum and a function of log diameters. This frequency can
Wise 1976; Neukum and Ivanov 1994; Neukum either be based upon the cumulative crater diame-
et al. 2001a; Strom et al. 2005). ter sizes (cumulative plots) in which the number of
Lunar-like cratering chronology models were craters with diameters equal or greater than D: N =
derived for Mercury (Strom and Neukum 1988; N(D) is presented or in differential form where
Neukum et al. 2001b), ▶ Venus (McKinnon the derivative dN/dD of the cumulative size-
et al. 1997), and Mars (Neukum and Wise 1976; frequency distribution gives the number of craters
Neukum and Hiller 1981; Hartmann and Neukum in equal diameter bins. The incremental form pro-
2001). Crater size-frequency measurements were vides the frequency as a function of the geometric
conducted also for a number of asteroids, e.g., average of fixed-diameter increments (usually √2).
951 Gaspra (Neukum and Ivanov 1994; Chapman The relative distribution (R-plot) is the deviation
et al. 1996), 243 Ida (Neukum and Ivanov 1994), of the size-frequency distribution from a power
and 253 Mathilde (Chapman et al. 1998). law: R = D3 (dN/dD). The choice depends on the
The impact-cratering chronology model for details that need to be depicted and highlighted
the Moon is characterized by an exponentially and the analyses that are carried out subsequently.
declining impact and crater formation rate in the The size-frequency distribution of an impact
first 1 Ga following planetary formation 4.55 Ga crater population can be approximated using
(e.g., Neukum 1977; Neukum and Ivanov 1994). either three stepwise power-law segments
Since about 3.8 Ga ago, impact and cratering (Hartmann et al. 2000) obtaining coefficients for
rates have dropped considerably and reached a the approximation that are characteristic for each
more or less constant level at 3–3.3 Ga ago SFD. In contrast to the approach by Hartmann,
(Wetherill 1975; Neukum 1983; Neukum and Neukum proposed an analytical polynomial fit
Ivanov 1994, Neukum et al. 2001a). Some authors (Neukum 1983; Neukum and Ivanov 1994;
interpreted a peak in radiometric ages of lunar Ivanov 2001; Neukum et al. 2001a). These func-
rocks at about 3.9 Ga as an indication for a strong tions are termed production functions and are
peak in impact and cratering rate (e.g., Tera characteristic of each planetary body and derived
et al. 1974). They concluded that this so-called from numerous observations and measurements
Late Heavy Bombardment (LHB) was of impact craters. Once the production function is
fitted, the frequency of impact craters larger than and many preliminary results are expected to be
a given diameter (usually N for D 1 or N for improved in the course of ongoing planetary
D 10 km) is obtained. This frequency value is missions.
subsequently used to derive a surface age from
the chronology function obtained earlier which The Moon
represents the impact-cratering record in the The Earth and its Moon are the only bodies in the
planet’s history. Solar System for which radiometric ages of rock C
Stratigraphic schemes have been established surface units and soil samples have been
in principle for all terrestrial bodies. These obtained. Robotic and manned missions to the
schemes are based upon stratigraphic marker Moon have provided the most extensive data set
horizons that are defined by geologic criteria concerning stratigraphy and surface ages avail-
which are different for each object, and they able (Wilhelms 1987).
take into account the superimposed crater size The lunar geologic history is subdivided into
frequencies considered to be a characteristic of periods defined by impact events and impact cra-
a particular unit (Wilhelms 1987). The geologic ter occurrences ranging from the pre-Nectarian
history of each planet is subdivided into the (oldest period), Nectarian, Imbrian,
so-called time-stratigraphic systems. Such Eratosthenian to the Copernican period
chronostratigraphic divisions are termed systems (youngest). The Imbrian period is subdivided by
and are further divided into several (e.g., lower, the Orientale impact into a late (upper) and early
middle, or upper) series. Time-stratigraphic sys- (lower) Imbrian epoch (Wilhelms 1987).
tems and series correspond to periods and epochs
as chronologic or chronometric divisions • The pre-Nectarian covers that section of geo-
(Wilhelms 1987). The beginning of each period logic history which predates the Nectarian
or epoch is defined by the cratering model age period, i.e., older than 3.92 (Wilhelms 1987;
derived from the SFD measurement on the unit Stöffler and Ryder 2001) to 4.1 Ga (Neukum
which defines the base of each system. and Ivanov 1994). The chronostratigraphic
system comprises a number of 30 impact
basins, among which is the South Pole-Aitken
Key Research Findings basin. No traces for volcanic or tectonic pro-
cesses during the pre-Nectarian period have
Stratigraphic sequences have been established in been found thus far. Traces of pre-Nectarian
principle for all terrestrial bodies in the Solar surface units are mainly found on the lunar
System, in particular for the Inner Solar System farside, and samples returned during the
planets and the Moon. The main stratigraphic Apollo and Luna missions are later-generation
sequences and unit-forming events are summa- material reworked as breccias in the course of
rized below, and ages are provided whenever subsequent impact processes (Wilhelms 1987;
possible. However, age uncertainties are Stöffler et al. 2006).
excluded for all model ages, and the reader is • The Nectarian period is defined as the time
referred to references listed in each section. period between the formation of the Nectaris
While for several bodies, such as the Moon and basin (Janssen Formation) and the Imbrium
Mars, chronostratigraphic systems are well impact event and comprises at least 11 other
established and applicable, other objects are large impact-basin forming events. Although
partly still unknown when it comes to high- some Nectarian-aged volcanic material has
resolution data needed for cratering analysis and been observed (Wilhelms 1987), much of
statistics. Consequently, differences in absolute this unit is covered by later impact events
age dating for each planet between various that masked Nectarian units. According to
groups of investigators are due to differences in recent studies, mare volcanism started already
assumptions of the impact and cratering rates, during the Nectarian period with ages of
3.92 Ga and continuing up to 1.2 Ga • The Copernican period is defined through the
(Hiesinger et al. 2003). The chronostra- occurrence of rayed craters on the Moon with
tigraphic basis, i.e., the lowermost unit, for Copernicus being the most prominent rayed
the Nectarian system is set between 3.92 impact crater. These impact craters are
(Wilhelms 1987; Stöffler and Ryder 2001) superimposed on all other units and show
and 4.1 Ga (Neukum and Ivanov 1994). traces of ejecta material all over the Moon
• The Imbrian period is defined by the impact- despite their relatively small size. Due to its
basin event forming the Imbrium impact crater definition, the basis for the Copernican period
on the lunar nearside with the Fra Mauro For- is not well constrained with ages ranging from
mation at its base. The Late Imbrian epoch 1.1 (Wilhelms 1987) to 1.5 Ga (Neukum and
starts with the Orientale basin formation Ivanov 1994; Neukum et al. 2001a) up to
(Hevelius Formation); its upper limit is defined 1.1–2.1 Ga (Stöffler and Ryder 2001). The
through impact crater sizes only. Two thirds of Copernican period continues up to the
the lunar mare volcanic deposits are of Late present time.
Imbrian age. Apart from extensive mare-type
volcanism, dark mantling deposits are thought Mercury
to be of Late Imbrian age (Stöffler et al. 2006). Mercury’s surface is visually comparable to the
Formation of the Orientale basin is suggested to lunar one as expressed by numerous impact cra-
be 3.72 Ga (Stöffler and Ryder 2001) to 3.8 Ga ters and large impact basins, as well as a rich
(Wilhelms 1987) or 3.84 Ga (Neukum and variety of tectonic features probably caused by
Ivanov 1994; Neukum et al. 2001a) depending rapid cooling and tidal despinning (Spudis and
on the chronology model. Guest 1988; Neukum et al. 2001a). Mercury’s
The Early Imbrian epoch covers the time surface shows a global dichotomy in terms of
range between Imbrium (Fra Mauro Forma- major geologic units: the densely cratered terrain
tion) and Orientale basin formation (Hevelius (highlands) with interspersed smoother areas
Formation) corresponding to 3.77 Ga (Stöffler (intercrater plains) and the less-densely cratered
and Ryder 2001) to 3.84 Ga (Wilhelms 1987) lowland plains (smooth plains) (Trask and Guest
or even 3.92 Ga (Neukum and Ivanov 1994; 1975; Spudis and Guest 1988). In contrast to the
Neukum et al. 2001a) depending on the chro- lunar surface, the smooth plains and highland
nology model. The epoch is characterized by terrain are comparable in relative albedo. Not-
extensive volcanism and impact cratering but withstanding the surficial resemblance of Mer-
without formation of larger impact basins, cury and the Moon, crater size frequencies of
except for the Schrödinger basin. Many of the heavily cratered highland terrain of Mercury
the light plains have a Lower Imbrian age are less than that of the Moon (Spudis and Guest
(Stöffler et al. 2006). 1988) with a general paucity of impact craters in
• The Eratosthenian period is defined through the 30 km diameter range (Neukum et al. 2001b).
the appearance of younger impact craters cov- The stratigraphic system of Mercury was
ering most of other lunar units but showing no established by geologic mapping at scales of
signs of rayed ejecta – a criterion that has led 1:5,000,000 in the late 1970s and early 1980s on
to discussions regarding boundary ambiguities image data of Mariner 10 flybys.
(Stöffler et al. 2006; Stöffler and Ryder 2001). Mercury’s surface resemblance to the Moon
Some of the mare basalts were emplaced dur- led to establishing a stratigraphic system compa-
ing the Eratosthenian, but these units are much rable to that of the lunar one with impact events
less extensive than those in the Imbrian characterizing bases of stratigraphic periods.
period. The basis of the Eratosthenian is gen- Mercury’s geologic history is therefore
erally set to 3.2 Ga (Wilhelms 1987; Stöffler subdivided into five periods starting with the
and Ryder 2001; Neukum and Ivanov 1994; pre-Tolstojan as the oldest unit to the Kuiperian
Neukum et al. 2001a). as the youngest period (Spudis and Guest 1988).
• The pre-Tolstojan period closely compares to 1988; Neukum et al. 2001a). As for the Moon,
the lunar pre-Nectarian and encompasses geo- the period is predominantly characterized by
logic units older than 3.97 Ga (Neukum young impact crater materials.
et al. 2001a) to 4.1 Ga (Strom and Neukum
1988) and is related to crater materials and Venus
multiring basins as well as intercrater plains Radar mapping investigations of Venus’ surface
(Neukum et al. 2001a; Tanaka and Hartmann in the context of the 1990s Magellan mission C
2008; Spudis and Guest 1988). have shown that the impact crater frequency on
• During the Tolstojan period which is equiva- Venus is extremely low. This indicates that the
lent to the lunar Nectarian period, the domi- surface as it is observed today has an age of few
nant geologic units are the Goya formation 100 Ma only (Basilevsky and Head 1998). Wide-
that marks the basis of the Tolstojan system spread volcanism and intensive tectonic disrup-
and which hosts deposits of the Tolstoj basin tions have shaped the surface and deleted most of
as well as materials of small impact basins and the older geologic units and the planet’s impact
craters. Its base is estimated to be approxi- crater record. Consequently, the stratigraphic
mately 3.9–4.0 Ga (Spudis and Guest 1988) system is not constrained throughout all of
up to 4.06 Ga (Strom and Neukum 1988) old. Venus’ history and could not be established
Recent age estimates based on new chronol- until the late 1990s, on the basis of photogeologic
ogy models put the Tolstojan basis at 3.97 Ga interpretation of a few selected regions that
ago (Neukum et al. 2001b). proved to be statistically significant for
• The Calorian period compares to the lunar establishing a global stratigraphic system
Imbrian period and is mainly characterized (Basilevsky and Head 1998).
by Caloris-group units, i.e., mountain mate- The current stratigraphic record represents only
rial, intermontane plains, hummocky plain, 10–20 % of Venus’ history and covers only 30 %
the Calorian plains, as well as impact crater of Venus’ surface as mapped at scales of
materials and materials of small impact 1:3–1:10 M (Basilevsky and Head 1998). Specific
basins. Its time-stratigraphic base is estimated geologic time units (periods) are defined mainly
to be in the range of 3.77 Ga (Neukum on the basis of large impact events and volcanic
et al. 2001a), 3.85 Ga (Strom and Neukum plain formation events, which helped to establish
1988) to 3.9 Ga (Spudis and Guest 1988) and four major periods: the pre-Fortunian (oldest),
is defined by the Caloris impact event that is Fortunian, Guineverian, and Aurelian (youngest
considered to be the youngest impact basin on unit). Due to the paucity of impact craters and
Mercury (McCauley et al. 1981). statistically relevant average estimates of surface
• The Mansurian period is equivalent to the ages, absolute ages are usually provided as ratios
lunar Eratosthenian period, and its chronostra- with respect to the global impact crater frequency
tigraphic base is defined by the impact event of (T = 1.98 106 craters/km2) corresponding to
Mansur. With some uncertainties, its crater an average surface age of 288 Ma (+311/98 Ma)
model age is estimated at 3–3.5 Ga (Spudis according to Strom et al. (1994), 400–800 Ma
and Guest 1988; Strom and Neukum 1988, according to Phillips et al. (1992), or as high as
Neukum et al. 2001a). As for the Kuiperian 800 Ma (+800/400) according to Zahnle and
system, major units are of impact crater origin McKinnon (1996) as discussed in Basilevsky and
mainly. Head (1998) in detail.
• The Kuiperian period represents the youngest
period in Mercury’s history, and its base is • The pre-Fortunian period is not constrained in
defined by the age of impact crater Kuiper terms of geologic surface units and spans the
which occurred 1 Ga ago, equivalent to the time before 1.47 T.
lunar Copernican system (Wilhelms 1987; • The Fortunian period is characterized by
Spudis and Guest 1988; Strom and Neukum intensive tectonic deformation with formation
of the ancient ▶ tessera terrain that covers plains. The subdivision of the stratigraphic system
about 8 % of Venus’ surface. Tesserae forma- of Mars is based on marker horizons that are
tion occurred at 1.47 T (1.93–1.01 T in previ- formed by plain-forming volcanism (Scott and
ous estimates) which is considered to form the Carr 1978; Tanaka 1986; Tanaka et al. 1992). Mar-
time basis of the Fortunian period (Basilevsky tian geologic time periods are from oldest to youn-
and Head 1998). gest: the Noachian (with late, middle, and early
• During the Guineverian period, extensive vol- epochs), the Hesperian (with a late and early
canic plains were formed with a peak at 1.1 T epoch), and the Amazonian (with a late, middle,
(Tanaka et al. 1997). The period has been pro- and early epoch) (Scott and Carr 1978; Tanaka
posed to be a supergroup consisting of four 1986; Tanaka et al. 1992). For Mars, several chro-
plain-forming subunits by Basilevsky and nology models were proposed and modified in the
Head (1998). These groups (youngest to oldest course of the availability of new higher-resolution
unit) follow the proposal by Basilevsky and data. Some of these efforts have been combined
Head (1998): the Atla Group consisting of rel- lately to form the recent chronology model by
atively undisturbed mafic lava, the Rusalka Hartmann and Neukum (2001). Stratigraphic
Group lava materials covering up to 75 % of boundaries (boundaries of time periods) are
Venus’ surface, the Lavinia Group character- slightly different, which has led to a Hartmann
ized by ridged and fractured plains, and the model (HM) and a Neukum model (NM).
Sigrun Group consisting of densely fractured
plain material emplaced as mafic lavas. The • A pre-Noachian period is informally
Guineverian period covers most of Venus’ established although the Noachian basis is
known geologic record and terminates at not exposed; however, radiometrically
around 0.1–0.2 T according to most recent esti- derived ages for the Martian meteorite
mates (Basilevsky and Head 1998). A period of ALH84001 with a crystallization age of
younger volcanic activity peaking at 0.4–0.5 T 4.5 Gyr fit into this period (Mittlefehldt 1994).
ago with formation of Venus’ coronae and rifts • The Noachian system is characterized by the
as well as formation of large volcanoes at 0.3 T oldest, densely cratered units in the highlands
was proposed as a fourth major unit in Tanaka covering a time range of older than 3.97
et al. (1997). –3.74 Ga for the Noachian system (Tanaka
• The Aurelian period is characterized by mate- et al. 1992; Hartmann and Neukum 2001). Its
rials associated with the youngest impact cra- basis is defined by highland material of the
ters; its basis is set to 0.1 T containing Noachis Terra located between the Argyre
approximately 10 % of Venus’ visible and Hellas Planitia impact basins (Tanaka
cratering record (Tanaka et al. 1997 and refer- et al. 1992). The system is generally charac-
ences therein) and translating to an age of terized by Heavy Bombardment impacts,
50 Ma using a conservative estimate of the large-scale volcanism peaking in the Tharsis
mean surface age. region and the highland volcanic provinces,
global tectonism and extensive valley network
Mars formation indicating fluvial processes, and a
On Mars, impact cratering and plain volcanism much denser atmosphere and warmer climate.
played a dominant role in shaping the planet’s The Late Noachian period spans
surface throughout history. Additionally, a rich 3.86–3.74 Ga (Hartmann and Neukum 2001)
variety of processes related to fluvial, glacial, and and is characterized by cratered plateau mate-
eolian resurfacing have significantly contributed to rial mainly. During that period, the crustal
the morphologies that are observed nowadays. dichotomy has been morphologically shaped.
Mars’ global topography is divided into the densely The Middle Noachian epoch which covers the
cratered and old highland terrain in the south and time span between 3.97 and 3.86 Ga (Hartmann
the smooth, less-densely cratered younger northern and Neukum 2001) is characterized by cratered
highland terrain shaped by impact cratering (NM). The period is generally defined through
and the Argyre Planitia impact-basin event. processes related to the northern lowland units
During the late Early Noachian (>3.97 Ga), and plain materials and is characterized by
the Hellas and Isidis impact basins formed, extensive resurfacing processes. Late-stage
and global volcanism shaped the Tharsis region volcanism and eolian resurfacing shaped
with the formation of Paterae and Tholi and the large areas of Mars and obliterated older
circum-Hellas Planitia Highland volcanoes units (Tanaka et al. 1992). The Amazonian C
(Tanaka et al. 1992). Recently, the Noachian also shows late-stage outflow activity in the
period was characterized in terms of geochem- circum-Chryse Planitia area and an abundance
ical alteration by Bibring et al.( 2006), of ice-related surface processes predominantly
suggesting that formation of clay minerals, near the global dichotomy escarpment and
i.e., phyllosilicates, peaked during the Early circum-Tharsis volcanoes as well as the
and Middle Noachian epochs. circum-Hellas/Argyre Planitia regions. Sur-
• The Hesperian period is subdivided into the face alteration by formation of anhydrous fer-
Early and Late Hesperian epochs spanning ric oxides led to the planet’s characteristic red
3.74–2.9 Ga and is characterized by the Hes- surface color (Bibring et al. 2006).
peria Planum ridged plain material northeast The basis of the Upper Amazonian series
of the Hellas Planitia impact basin. Impact- (Late Amazonian epoch) is defined by flood
cratering rates were significantly lower when plain material of the southern Elysium Planitia
compared to the Noachian period, marking the area and starts 0.3 Ga (HM) to 0.6 Ga
end of the Heavy Bombardment period. (NM) ago (Tanaka 1986; Hartmann and
Vanishing fluvial activity was replaced by Neukum 2001). Characteristic materials of
large-scale volcanism in the lowland units the Upper Amazonian are predominantly
(Tanaka et al. 1992 and ref’s therein). The found near the seasonally changing polar
disappearance of surface water has led to the caps of Mars and in the young deposits and
assumption that most of the water is stored as flow fields delineating the northern hemi-
permafrost under the surface. Catastrophic spheric volcanoes. The Middle Amazonian is
release of water led to formation of outflow defined by the age of Amazonis Planitia lava
channels on Mars in the circum-Chryse and flow materials (Tanaka 1986; Hartmann and
eastern Hellas Planitia regions along with for- Neukum 2001) and covers the time range
mation of the Martian chaotic terrain and the between 2.1 and 0.6 Ga (NM) or 1.4–0.3 Ga
Valles Marineris system (Tanaka et al. 1992). (HM). The basis for the Early Amazonian
The Hesperian is also characterized by exten- epoch is set to 3.3 Ga (NM) to 2.9 Ga
sive sulfate deposits (Bibring et al. 2006) pri- (HM) ago and is defined by the smooth plain
marily in the Valles Marineris region. The materials of Acidalia Planitia (Tanaka 1986;
Early Hesperian period covers the age range Hartmann and Neukum 2001).
of 3.74–3.65 Ga (Hartmann and Neukum
2001) and is defined through the Hesperia Outer Solar System Objects
Planum ridged plain units. Geologic materials In contrast to the currently accepted chronology
of the Late Hesperian, covering the period models proposed for the inner planets which are
between 3.65 and 2.9 Ga (Hartmann and based upon the lunar chronology model with
Neukum 2001), are defined through the plain impactors from the asteroid belt, planetary
material of the northern plain Vastitas objects in the Outer Solar System are treated in
Borealis unit. a different way. While one group favors the
• The Amazonian period spans much of Martian lunar-like distribution and chronology model for
history and starts 2.9 Ga ago according to the all objects in the Solar System, i.e., lunar-like
Hartmann model (HM) and up to 3.31 Ga ago time dependence of the cratering rate, the second
according to the Neukum chronology model group favors nearly constant cratering rates by
cometary impactors. A number of researchers chronology model. However, resurfaced units, in

developed lunar-like chronology models for the particular on Enceladus and Dione, show lower
Jovian and Saturnian system (Shoemaker and ages at approximately 1.5 Ga (Boyce and Plescia
Wolfe 1982; Boyce and Plescia 1985; Neukum 1985; Neukum 1985).
1985, 1997). Based on present-day estimates of Current efforts focus on updating existing
sizes and dynamics, a constant cratering rate chronology models with the help of new space-
chronology for the icy satellites was put forward craft data (e.g., Zahnle 2003) and imply ages far
with impactors originating in the Kuiper Belt below 1 Ga for the resurfaced terrain on
(Zahnle et al. 1998). Enceladus and ages as old as 4 Ga for the
The two competing cratering chronology older terrain, using a constant-rate chronology
models agree well for old, densely cratered sur- model with mainly cometary contribution of
faces on the icy satellites, but they are different impactors. Mimas’ surface crater impact density
by more than an order of magnitude for younger, is close to saturation, suggestive of an old age,
resurfaced units. These issues are discussed in but it lacks larger impact craters beyond 30 km
detail in, e.g., Neukum (1997, 1998) and Zahnle implying a much younger surface age. For Sat-
et al. (1998). urn’s moon Titan, the small number of observed
Imaging data of the Voyager and Galileo mis- impact craters suggests very young surface ages.
sions to the Outer Solar System provided the A closer look at the data and impact conditions,
basis for impact crater size-frequency statistics however, shows that much of Titan’s surface is
and estimates of surface ages of the Galilean probably as old as 2 Ga (Jaumann and Neukum
satellites. The surface of Europa is considered to 2009).
be relatively young under assumption of primar-
ily cometary impacts with average ages in the
range of 30–70 Ma, as suggested by the less-
See Also
densely cratered surface of Europa when com-
pared to Callisto or Ganymede (Lucchitta and
▶ Crater, Impact
Soderblom 1982; Zahnle 2003). Employing a
▶ Mare, Maria
lunar-like chronology model with the asteroid
▶ Mars
belt as main impactor source, surface ages of
▶ Mercury
Europa are in the range of 0.5–1.5 Ga. However,
▶ Moon, The
individual units can be as young as approximately
200 Ma or less (Neukum et al. 1998).
▶ Tessera, Tesserae
The heavily cratered dark plains on Callisto ▶ Venus
and Ganymede are on the order of 4 Ga and older
in both asteroidal and cometary-source chronol-
ogy models (Neukum et al. 1998; Zahnle References and Further Reading
et al. 1998, Zahnle 2003). For the younger
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Fulchignoni M, Moore H, Neukum G, Schultz P,
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Circumbinary Planet 467
Definition
Planet
In chemistry, circular dichroism (CD) refers to
the differential absorption of left- and right- r P1-Sec
handed circularly polarized light by optically
active chiral molecules. Electromagnetic radia- Secondary r P1-Pr
tion consists of electric and magnetic fields that C
oscillate perpendicular to one another and to the CoM
direction of propagation. Linearly polarized light

occurs when the electric field vector oscillates
only in one plane and changes in magnitude, Primary
while circularly polarized light occurs when the
electric field vector rotates about its propagation
direction and retains constant magnitude. It thus
forms a helix propagating in space.
When circularly polarized light passes through
an optically active light-absorbing medium, the Planet
velocities of its right and left polarizations differ
as does the extent to which they are absorbed. CD r P1-Sec
is a measure of the difference in the amount of
absorption between the two. Since circularly Secondary r P1-Pr
polarized light is “chiral,” it interacts differently
with the two enantiomers of chiral molecules. f-q
rSec-Pr q
Depending on the wavelength employed, CD can
be used to investigate the structures of proteins, Primary
nucleic acids, small organic molecules, and
charge-transfer transitions.
See Also
▶ Chirality
▶ Polarized Light and Homochirality Circumbinary Planet, Fig. 1 General schematic presen-
tation of a P-type system. The top panel shows the general
orbital configuration of the planet and the binary stars.
Both stars and the planet rotate around the center of
mass of the binary (shown by CoM). However, it is cus-
tomary to consider the primary star to be stationary and
Circumbinary Planet both the secondary and planet rotate around this star
(bottom panel) (Figures from Haghighipour and
Kaltenegger (2013))
Nader Haghighipour
in a binary sar system. These orbits have been
historically labeled as:
Definition
1. The S-type (or Satellite-type) orbit where the
From dynamical point of view, three types of planet revolves around one of the stars of the
orbits are recognized for the motion of a planet binary (see ▶ circumprimary planet).
468 Circumplanetary Disk
2. The P-type (or Planetary-type) orbit where the

planet revolves around the entire binary sys- Circumprimary Planet
tem (circumbinary orbit), and
3. The L-type (or Libration-type) where the Nader Haghighipour
planet moves in the same orbit as the second- Institute for Astronomy, University of Hawaii-
ary star, but 60 ahead or behind it. Manoa, Honolulu, Hawaii, HI, USA
Figure 1 shows a circumbinary (P-type) orbit.

In general, the planet and the binary revolve Definition
around their common center of mass. However,
it is customary to consider the primary star to be From a dynamical point of view, three types of
stationary and both, the secondary and planet orbits are recognized for the motion of a planet in
rotate around it. a binary star system. These orbits have been
Although observations of disks in circumbinary historically labeled as:
orbits had indicated that planets must exist in these
orbital configurations, it was not until 2011 that 1. The S-type (or satellite-type) orbit where the
using the data from the Kepler Space Telescope, a planet revolves around one of the stars of the
team of scientists lead by Laurance Doyle discov- binary (see circumprimary planet)
ered the first circumbinary planet (Kepler 16b). 2. The P-type (or planetary-type) orbit where the
During the next two years, scientists discovered planet revolves around the entire binary sys-
many more circumbinary planets using the Kepler tem (circumbinary orbit)
telescope (see references below) 3. The L-type (or libration-type) where the
planet moves in the same orbit as the second-
ary star, but 60 ahead or behind it
References and Further Reading Figure 1 shows a circumprimary (S-type)

orbit. In general, the planet and the binary revolve
Doyle LR, Carter JA, Fabrycky DC et al (2011) Science around their common center of mass. However, it
333:1602
is customary to consider the primary star to be
Haghighipour N, Kaltenegger L (2013) Astrophys
J 777, article id. 166 stationary, and both the secondary and planet
Kosov VB et al (2014) Astrophys J 787, article id. 93 rotate around it.
Kostov VB et al (2013) Astrophys J 770, article id. 52 Many circumprimary planets have been dis-
Kostov VB et al (2014) Astrophys J 784, article id. 14
covered during the past 20 years. A complete and
Orosz JA, Welsh WF, Carter JA et al (2012a) Astrophys
J 758:87 up-to-date list of these planets can be found at
Orosz JA, Welsh WF, Carter JA et al (2012b) Science http://www.univie.ac.at/adg/schwarz/multiple.
337:1511 html.
Schwamb ME, Orosz JA, Carter JA et al (2013) Astrophys
Almost all these binary systems are wide with
J 768:127
Welsh WF, Orosz JA, Carter JA et al (2012) Nature separations between 250 and 6,000 AU. There
481:475 are, however, six circumprimary systems in
which the binary stars have small separations
(20 AU): GL 86, g Cephei, HD 41004, HD
196885, HD 176051, and a Centauri (note that
Circumplanetary Disk the existence of the planet around the secondary
star of this system has been challenged; see
▶ Planetary Rings Alpha Centauri Bb). The formation of these
Circumprimary Planet 469
Circumprimary Planet,
Fig. 1 Schematic
presentation of an S-type
system. The two stars of the
binary, (primary, and
secondary) revolve around
their center of mass (CoM),
while the planet orbits only C
one of the stars (top panel).
It is, however, customary to
neglect the motion of the
binary around its CoM and
consider the motion of the
secondary around a
stationary primary (bottom
panel). (Figure from
Kaltenegger and
Haghighipour (2013))
planets in such tight binaries is one of the extrasolar planets, vol 294, ASP Conf. Ser. ASP, San
most outstanding questions in theoretical Francisco, p 43
Endl M, Cochran WD, Hatzes AP, Wittenmyer RA
exoplanetary science. See the book Planets in (2011) In: Schuh S, Drechsel H, Heber U (eds) Plane-
Binary Star Systems by Haghighipour (2010, tary systems beyond the main sequence, vol 1331, AIP
Springer) for a comprehensive and detailed Conf. Proc. AIP, Melville, p 88
review. Haghighipour N, Dvorak R, Pilat-Lohinger E (2010) In:
Haghighipour N (ed) Planets in binary star systems.
Springer, New York, p 539
Hatzes AP (2013) Astrophys J 770:133
Kaltenegger L, Haghighipour N (2013) Astrophys
References and Further Reading J 777, 165
Muterspaugh MW, Lane BF, Kulkarni SR et al (2010)
Correia ACM, Udry S, Mayor M et al (2008) Astron Astron J 140:1657
Astrophys 479:271 Queloz D, Mayor M, Weber L et al (2000) Astron
Dumusque X, Pepe F, Lovis C et al (2012) Nature 491:207 Astrophys 354:99
Eggenberger A, Udry S, Mayor M (2003) In: Deming D, Zucker S, Mazeh T, Santos NC, Udry S, Mayor M (2004)
Seager S (eds) Scientific frontiers in research on Astron Astrophys 426:695
470 Circumstellar Chemistry
asymptotic giant branch (AGB) phase. These

Circumstellar Chemistry stellar ejecta might be contaminated to a large
extent by material processed through stellar
Marcelino Agundez nucleosynthesis in the interior of the star, and
Instituto de Ciencia de Materiales de Madrid, thus the elemental composition may be quite
Madrid, Spain different from that of the primordial cloud from
which the star formed, with abundance enhance-
ments of some elements heavier than helium,
Keywords such as nitrogen and carbon (Iben and Renzini
1983; Woosley and Weaver 1986). Stellar mass
Astrochemistry; Circumstellar envelopes; Stellar loss processes give rise to circumstellar enve-
evolution; Chemical reactions; Interstellar dust; lopes, which are sites of an extraordinary interest
Mass loss; Molecular lines; Molecules; Sub- from a chemical point of view, as they act as
millimeter observations; Thermochemical factories of a population of tiny dust grains and
equilibrium a rich variety of molecules. In fact, most circum-
stellar envelopes are made up of molecular gas
and dust.
Definition Stellar ejecta provide the environment where
dust grains are synthesized by condensation of
Circumstellar chemistry refers to the group of refractory elements, and the ejection process pro-
processes able to transform the chemical nature vides the mechanism to inject them into the inter-
of a cloud of gas and dust surrounding a star. This stellar medium. It is well known that dust is a
general definition encompasses both young and minor (about 1 % in mass) but ubiquitous com-
evolved stars out of the main sequence, those ponent of interstellar clouds and that it is to a
which may host an envelope with a large extent responsible for their physical and
non-negligible mass. Most authors, however, chemical state. Dust grains, for example, provide
use the term “circumstellar chemistry” to refer an efficient protection against external ultraviolet
to the chemical processes occurring in the ejecta radiation, and their solid surfaces can act as cat-
of an evolved star, rather than in a cloud feeding alysts in the formation of molecules, including
the formation of a star or a planetary system. This the most abundant interstellar molecule, H2. It is
entry focuses on this second more restricted def- currently thought that most of the interstellar dust
inition. For a description of the chemistry in is in fact synthesized and supplied by the ejecta of
clouds around protostars and young stars, see dying stars (Gehrz 1989).
Hot Cores; ▶ Hot Corinos; and Protoplanetary Stellar ejecta also provide a suitable environ-
Disk, Chemistry. ment to synthesize and host a rich variety of
molecules, some of them quite exotic by terres-
trial standards. Thus, in IRC +10216, a prototyp-
Overview ical circumstellar envelope around an AGB star,
the molecular inventory accounts for about half
During their last stages of evolution, stars expe- of all the molecules ever discovered in space
rience important mass loss processes, which (▶ Molecules in Space). In fact, molecules
serve to recycle the stellar material back to the make up most of the mass of envelopes around
▶ interstellar medium. Massive stars (≳8 solar low- and intermediate-mass evolved stars and are
masses) end with a sudden and violent explosion a very important constituent of envelopes around
in the form of a supernova, while low- and massive evolved stars such as the red supergiant
intermediate-mass stars (≲8 solar masses) lose VY Canis Majoris (Ziurys et al. 2007) and yellow
their mass through more gentle stellar winds, hypergiants (Quintana-Lacaci et al. 2007). In the
which are particularly intense during the case of ▶ supernova ejecta, molecules such as
Circumstellar Chemistry 471
CO and SiO might still be present at abundance ratio of carbon to oxygen is C/O <1,
non-negligible levels, as in the case of SN during the AGB phase convective (dredge up)
1987A (Petuchowski et al. 1989; Liu and processes inside the star may bring out to the
Dalgarno 1994), but in general the formation stellar surface an amount of carbon high enough
and survival of molecules is severely hampered to exceed that of oxygen (Iben and Renzini 1983).
by the harsh irradiation conditions to which these Therefore, depending on whether the elemental
ejecta are subjected. Molecules in circumstellar C/O abundance ratio at the stellar surface is lower C
envelopes around evolved stars are relatively or higher than 1, AGB stars may be either oxygen
short-lived in astronomical terms (usually less or carbon rich, respectively. At the temperatures
than a few tens of thousands of years), since on and pressures of the atmospheres of AGB stars,
their way from the star to the interstellar medium, thermochemical equilibrium governs the chemical
they become completely exposed to the ultravio- composition, which is essentially molecular,
let interstellar radiation field at the outer edge of although the relative abundance of different mol-
the envelope, where they are ultimately photodis- ecules will depend on the elemental C/O abun-
sociated. Therefore, unlike the case of dust, free dance ratio. The high stability of CO makes this
molecules are destroyed before being incorpo- molecule reach very high abundance, locking up
rated into the interstellar medium. most of the limiting element (either C or O) and
Mass loss processes in evolved stars enrich the allowing for the element in excess to form, either
interstellar medium with metals (which in astro- oxygen-bearing molecules such as H2O and SiO if
nomical terms refer to any element heavier than C/O <1, and the opposite, i.e., carbon-bearing
helium) and dust grains and provide the main molecules such as C2H2 and HCN, if C/O >1.
mechanism by means of which the interstellar This fact is firmly established by theoretical cal-
medium is recycled with fresh material. culations (Tsuji 1973) and has been also exten-
When matter is expelled from the stellar atmo- sively confirmed by observations. AGB stars have
sphere and expands into outer space, it experi- an spectral type M (effective temperature between
ences a wide range of physical conditions. The 2,000 and 3,700 K), but it is interesting to note that
number density of particles decreases by several distinct spectral signatures allow to classify them
orders of magnitude, and the temperature drops according to their elemental C/O abundance ratio
from thousands of Kelvin to just 10–20 K. Along into M stars (reserved for stars with C/O <1),
the outflow, matter is chemically processed by a C stars (with C/O >1), and S stars (with C/O
wide variety of processes, such as three-body and ~1). The chemical composition at the stellar sur-
endothermic chemical reactions in the hot surface propagates to the base of the envelope and, to
roundings of the atmosphere, condensation of a large extent, to the outer layers, and it is thus of
dust grains a few stellar radii away from the great importance for the overall chemical compo-
star, and photodissociation and photoionisation sition of the circumstellar envelope. In fact, obser-
in the outer layers, whose resulting radicals and vations of molecular lines arising from outer
ions take part in exothermic and rapid neutral- circumstellar layers (e.g., Bujarrabal et al. 1994)
neutral and ion-neutral chemical reactions. Here- have confirmed the existence of circumstellar
after, an overview is given on the chemical state envelopes with two different types of chemical
and chemical processes at work along the various composition, which correspond to the oxygen- or
regions traveled by matter in its journey from the carbon-rich character of the star.
star to the interstellar medium (see Fig. 1). The ultimate mechanism leading to the ejec-
tion of matter from a red giant star, and thus to an
Stellar Atmosphere and Inner Envelope effective mass loss, is not completely understood,
One of the most important parameters that define although it is very likely related to both waves
the chemistry of a circumstellar envelope is the induced by the pulsation of the star and radiation
elemental C/O abundance ratio at the photo- pressure on dust grains (Gilman 1972; Kwok
sphere of the central star. Although the cosmic 1975; Bowen 1988). Mass loss processes during
Circumstellar Chemistry, Fig. 1 Scheme of a circum- of condensation of dust, and rPH indicates the region
stellar envelope around an AGB star. The envelope is where molecules start to be photodissociated by the ambi-
divided into three regions: stellar atmosphere, inner enve- ent interstellar ultraviolet field. The bottom axes indicate
lope, and outer envelope. The top axis indicates various the typical temperatures T and number densities of parti-
characteristic radii from the star: R* is the stellar radius, r0 cles n at the various locations in the envelope
indicates the end of the stellar atmosphere, rc is the radius
the AGB phase are particularly intense, with material pushed out by the emerging wave
mass loss rates between 107 and 104 solar (Hinkle et al. 1982). On the other hand, a few
masses per year and expansion velocities around stellar radii away from the AGB star, once the
10–20 km s1 (Olofsson 2008). Whatever the kinetic temperature has fallen below about
exact mechanism of the mass loss, it occurs 1,000 K, those elements with a higher refractory
within a few stellar radii from the AGB star, a character begin to condensate out of the gas
region which is probably the one hosting the phase in the form of tiny dust grains. The chem-
most complex and not yet fully understood physical composition, originally established by ther-
ical and chemical processes that take place mochemical equilibrium at the stellar
throughout the whole envelope. On the one photosphere and surroundings, is then modified
hand, the periodic pulsation of the star gives a few stellar radii away from the star due to the
rise to shock waves which are formed between combined action of, at least, these two processes,
the gas infalling from the previous cycle and i.e., shock waves and dust formation.
There is evidence of deviations from thermo- to the possibility of observing high-excitation

chemical equilibrium in the chemical composi- molecular lines at mm- and submm wavelengths
tion of the inner regions of circumstellar with the unprecedented angular resolution
envelopes, which comes from the observation of of ALMA.
anomalously high abundances of HCN in A further interesting aspect of the chemistry of
oxygen-rich objects (Duari and Hatchell 2000; these inner parts of circumstellar envelopes is
Schöier et al. 2013), water vapor in carbon-rich that, due to the warm temperatures, a wide variety C
envelopes (Decin et al. 2010; Neufeld of refractory elements may survive in the gas
et al. 2011), and ammonia (NH3) in both oxygen- phase and participate in the formation of some
and carbon-rich envelopes (Keady and Ridgway exotic molecules. This is not the case for inter-
1993; Justannont et al. 2012). Recent results stellar clouds, where the cold temperatures make
obtained with the Herschel Space Observatory most refractory elements readily condense out of
have widely contributed to highlighting the the gas phase. Metals (in the chemical, not astro-
departure from thermochemical equilibrium of nomical, sense) such as Na, K, Al, and Ti take
inner layers of envelopes around evolved stars. part in the formation of molecules such as NaCl,
Endothermic and three-body chemical reactions AlCl, KCl, AlF, NaCN, AlO, AlOH, TiO, and
are effective at the high temperatures and densi- TiO2 (Cernicharo and Guélin 1987; Kamiński
ties of these inner layers and must play an impor- et al. 2013) in circumstellar envelopes. The
tant role in the departure from chemical observed abundances of these molecules are gen-
equilibrium. However, the ultimate mechanism erally in good agreement with the predictions of
triggering this nonequilibrium is currently not thermochemical equilibrium calculations.
well understood. Efforts have been made to
model the impact of shock waves on the chemical Outer Circumstellar Layers
composition of the inner envelope around an It is well established by interferometric maps of
AGB star (Cherchneff 2006, 2012; Agúndez and molecular line emission that molecules in cir-
Cernicharo 2006). It has been also suggested that cumstellar envelopes are either concentrated
the clumpy character of circumstellar envelopes around the central star or distributed in a hollow
may allow interstellar ultraviolet photons to pen- shell located far from the star. In the case of IRC
etrate into the inner regions and to induce a warm +10216, for example, molecules such as HCN,
photochemistry (Decin et al. 2010). Dust forma- CS, or SiS fall in the former category, while
tion is also likely to affect the abundances of gas others such as CN, HNC, HC3N, C2H, or C4H
phase molecules in these inner regions, on the one belong to the latter group (Guélin et al. 1997).
hand, because the formation of dust must occur at These two distributions are clearly denoting two
the expense of the gaseous material, and thus different origins for each group of molecules.
some molecules must decrease their abundances Those which are concentrated around the star
and, on the other, because chemical reactions are formed at chemical equilibrium in the stellar
may occur on the surface of grains resulting in atmosphere or through any of the nonequilibrium
an important chemical processing of the gas. An processes at work in the warm inner layers. On
adequate theoretical description of the formation the other hand, those molecules which appear at a
of dust is hampered by the fact that it probably certain radial distance from the star must origi-
occurs out of chemical equilibrium (Gail and nate by some other process, which seems to be
Sedlmayr 1988). Many uncertainties exist on the triggered by the penetration of ultraviolet radia-
interplay between the various physical and chem- tion from the ambient interstellar medium.
ical processes at work in the inner regions of Indeed, as matter expands from the star, it is
envelopes around AGB. In fact, this is perhaps geometrically diluted (the number density of par-
the area of circumstellar chemistry least well ticles falls with the square of the radial distance to
understood, so that it will certainly experience the star for an isotropic expansion), and mole-
significant advances in the future, in part thanks cules and dust become increasingly exposed to
the external ultraviolet field. It has been argued of these molecules benefits from the extremely
that the appearance of a good number of mole- large rate constants of radiative association
cules at about the same radial distance from the between the neutral counterpart (usually a radical
star in IRC +10216 may be related to their with a moderately large size and a high electron
photodesorption from dust grain surfaces, onto affinity) and an electron.
which they would have been formed or deposited It is quite remarkable to note that metals sur-
in advance (Guélin et al. 1993). vive to an important extent in the cool outer
An alternative mechanism, which is supported layers of circumstellar envelopes, in spite of
by quantitative modeling calculations by differ- their high refractory character, which tends to
ent authors, proposes that molecules distributed bring them out of the gas phase as solid conden-
in the cool outer layers are formed in situ by gas sates. Neutral and ionized atoms of Na, K, Ca, Cr,
phase chemical reactions, which are ultimately and Fe have been observed in the outer layers of
driven by the photodissociation, and to a lesser IRC +10216 (Mauron and Huggins 2010) with
extent by photoionisation of the parent molecules abundances that represent a few percent of the
injected into the expanding envelope from the total abundance of the element. Also, metal-
inner layers (Glassgold 1996; Millar et al. containing molecules of the family of the cya-
2000). A circumstellar envelope around an AGB nides, such as MgNC, MgCN, HMgNC, AlNC,
star may be, in fact, viewed as a dynamically and FeCN, have been observed in the outer layers
expanding photon dominated region (PDR). As an of some circumstellar envelopes (see e.g., Guélin
example to illustrate how the chemical synthesis et al. 1993). Unlike the case of NaCl and the other
works in this scenario, we may focus on the case of molecules formed at chemical equilibrium in the
carbon-rich envelopes, where the photodissocia- surroundings of the star, these molecules seem to
tion of the parent molecules C2H2 and HCN benefit from the survival of metal atoms in the gas
produces the radicals C2H and CN, respectively, phase and form in situ in the cool outer layers by
which in turn participate in a chain of exothermic nonequilibrium chemical processes (Petrie
and fast neutral-neutral reactions leading to the 1996).
growth of carbon chains, such as polyynes and The variety and complexity of the molecules
cyanopolyynes. A wide variety of polyynes and observed in carbon-rich circumstellar envelopes
cyanopolyynes, and their related radicals, have is significantly greater than that observed in
been observed in carbon-rich circumstellar enve- oxygen-rich sources. This is not very surprising
lopes, noticeably in IRC +10216, a source that taking into account the larger availability of car-
concentrates much of the observational and theo- bon, not locked up into CO, in carbon-rich
retical research on circumstellar chemistry sources and the extraordinary chemical proper-
(Cernicharo et al. 2000). Ion-neutral chemical reac- ties of this element, which can easily form carbon
tions occur also in the outer layers of IRC +10216, skeletons giving raise to a wide variety of com-
but their ability to take part in the synthesis of plex molecules. However, it should be noted that
complex molecules is limited by the relatively the large differences in terms of variety and com-
low ionization fraction in these regions (~107). plexity of molecules between oxygen- and
Among the families of molecules observed in carbon-rich envelopes may be to some extent an
circumstellar envelopes, the most recent has been observational bias, a consequence of the over-
that of molecular anions. No negatively charged whelming attention paid by observers to carbon-
molecule was known in the interstellar or circum- rich sources, in particular to IRC +10216. Our
stellar medium prior to 2006. To date, up to six view on the relatively poor molecular content of
different molecular anions (C4H, C6H, C8H, oxygen-rich sources might change to some extent
CN, C3N, and C5N) have been discovered in in the near future, thanks to the great improve-
the expanding gas of IRC +10216 (see Agúndez ment of the capabilities of radiotelescopes, with
et al. 2010 and references therein). The formation particular regard to their sensitivity and spectral
coverage, which allows astronomers to carry out The protoplanetary nebula CRL 2688, although
more sensitive molecular line surveys and which more evolved than IRC +10216, still shares many
may extend significantly the molecular inventory similarities with this latter object in its chemical
of oxygen-rich envelopes (see e.g., Ziurys composition, with high abundances of unsatu-
et al. 2007). rated carbon chains such as C2H, C4H, and
HC3N. The protoplanetary nebula CRL 618 is
Beyond the AGB Phase clearly in a more advanced stage than CRL C
The evolution of a red giant star into a white 2688, so that some important chemical differ-
dwarf brings drastic changes on the chemical ences can be appreciated, such as the relatively
composition of the surrounding nebula. The star high abundances of the ions HCO+ and N2H+ and
shrinks enormously (the stellar radius decreases oxygen-bearing molecules such as H2CO and
from a few astronomical units down to about the H2O. It is also worth noting that some abundance
radius of the Earth), but it becomes also much ratios such as HNC/HCN and HC3N/HCN
hotter (the effective temperature increases from increase in CRL 618 and aromatic molecules
2,000–3,000 K up to 8,000–40,000 K) injecting such as benzene appear. Most of these changes
an intense ultraviolet radiation field in the sur- are thought to be the result of the strong chemical
rounding medium. This energetic radiation pho- processing driven by the ultraviolet field of the
todissociates most circumstellar molecules and star (Cernicharo 2004). Finally, the planetary
ionizes the resulting atoms, leading to the forma- nebula NGC 7027 shows some clear chemical
tion of a planetary nebula (PN). The transition differences with respect to less evolved counter-
from the AGB to the PN phase occurs rapidly parts. The circumstellar gas contains new mole-
(in about 103 years). It is however possible to cules such as the cations CO+, CH+, and HCS+,
identify some objects in this transition stage, and in general, the chemical complexity, in the
which are known as protoplanetary nebulae. The sense of molecules with a high number of atoms,
characteristics of protoplanetary nebulae are in decreases noticeably.
between those of the AGB and PN phases (Kwok One of the latest findings relative to the chem-
1993), i.e., they still host a circumstellar envelope istry of post-AGB objects has been the discovery
of gas and dust with a high molecular content, of the fullerenes C60, C70, and C60+ in some
although the effects of the ultraviolet radiation planetary nebulae (e.g., Cami et al. 2010).
field which emanates from the central star begin Indeed, it seems that there exists a clear differ-
to be visible. For example, there is an important ence between the chemistry of circumstellar
amount of ionized gas, and the spectra show a envelopes around AGB stars and post-AGB
series of unidentified infrared bands, which are objects, with regard to the appearance of some
usually assigned to polycyclic aromatic hydro- large ring and cluster carbon-based molecules,
carbons (PAHs). Another important distinctive such as benzene, PAHs, and fullerenes, in late
sign of protoplanetary nebulae is the appearance evolutionary stages. The formation of such
of fast bipolar winds (with velocities of hundreds molecular structures in these environments is
of km s1). yet to be understood.
The chemical processing to which circumstel-
lar envelopes are subjected during the evolution
from the AGB to the PN phase can be appreciated See Also
by comparing the molecular inventory in various
well-known carbon-rich objects in different evo- ▶ Asymptotic Giant Branch Star
lutionary stages, such as IRC +10216 (still in the ▶ Interstellar Chemical Processes
AGB phase), CRL 2688 (a young protoplanetary ▶ Interstellar Dust
nebula), CRL 618 (a protoplanetary nebula in a ▶ Molecules in Space
later stage), and NCG 7027 (a planetary nebula). ▶ Nucleation of Dust Grains
▶ Organic Dust, Influence on the Origin of Life Gilman RC (1972) On the coupling of grains to the gas in
▶ Organic Dust, Synthesis by Stars circumstellar envelopes. Astrophys J 178:423–426
Glassgold AE (1996) Circumstellar photochemistry. Annu
▶ Photochemistry Rev Astron Astrophys 34:241–277
▶ Photon Dominated Region Guélin M, Lucas R, Cernicharo J (1993) MgNC and the
▶ Planetary Nebula carbon-chain radicals in IRC +10216. Astron
▶ Protoplanetary Disk, Chemistry Astrophys 280:L19–L22
Guélin M, Lucas R, Neri R (1997) Mass loss in AGB stars.
▶ Protoplanetary Nebula IAUS 170:359–366
▶ Red Giant Hinkle KH, Hall DN, Ridgway ST (1982) Time series
▶ Star Dust infrared spectroscopy of the Mira variable star cCygni.
▶ Thermochemical Equilibrium Astrophys J 252:697–714
Iben I Jr, Renzini A (1983) Asymptotic giant branch
evolution and beyond. Ann Rev Astron Astrophys
21:271–342
References and Further Reading Justannont K, Khouri T, Maercker M et al (2012)
Herschel/HIFI observations of O-rich AGB stars:
Agúndez M, Cernicharo J (2006) Oxygen chemistry in the molecular inventory. Astron Astrophys 537:A144
circumstellar envelope of the carbon-rich star IRC Kamiński T, Gottlieb CA, Schmidt MR et al (2013)
+10216. Astrophys J 650:374–393 Dust-forming molecules in VY Canis Majoris (and
Agúndez M, Cernicharo J, Guélin M et al (2010) Astro- Betelgeuse). EAS Publ Ser 60:191–198
nomical identification of CN, the smallest observed Keady JJ, Ridgway ST (1993) The IRC +10216 circum-
molecular anion. Astron Astrophys 517:L2 stellar envelope. III – infrared molecular line profiles.
Bowen GH (1988) Dynamical modelling of long-period Astrophys J 406:199–214
variable star atmospheres. Astrophys J 329:299–317 Kwok S (1975) Radiation pressure on grains as a mecha-
Bujarrabal V, Fuente A, Omont A (1994) Molecular nism for mass loss in red giants. Astrophys
observations of O- and C-rich circumstellar envelopes. J 198:583–591
Astron Astrophys 285:247–271 Kwok S (1993) Proto-planetary nebulae. Annu Rev Astron
Cami J, Bernard-Salas J, Peeters E, Malek SE Astrophys 31:63–92
(2010) Detection of C60 and C70 in a young planetary Liu W, Dalgarno A (1994) Silicon monoxide in SN 1987-
nebula. Science 329:1180 A. Astrophys J 428:769–776
Cernicharo J (2004) The polymerization of acetylene, Mauron N, Huggins PJ (2010) Gas phase atomic metals in
hydrogen cyanide, and carbon chains in the neutral the circumstellar envelope of IRC +10216. Astron
layers of carbon-rich proto-planetary nebulae. Astrophys 513:A31
Astrophys J Lett 608:L41–L44 Millar TJ, Herbst E, Bettens RPA (2000) Large molecules
Cernicharo J, Guélin M (1987) Metals in IRC in the envelope surrounding IRC +10216. MNRAS
+10216 – Detection of NaCl, AlCl, and KCl, and 316:195–203
tentatively AlF. Astron Astrophys 183:L10–L12 Neufeld DA, González-Alfonso E, Melnick G et al (2011)
Cernicharo J, Guélin M, Kahane C (2000) A 12 mm The widespread occurrence of water vapor in the cir-
molecular line survey of the C-star envelope IRC cumstellar envelopes of carbon-rich Asymptotic Giant
+10216. Astron Astrophys Suppl 142:181–215 Branch stars: first results from a survey with Herschel/
Cherchneff I (2006) A chemical study of the inner winds HIFI. Astrophys J Lett 727:L29
of asymptotic giant branch stars. Astron Astrophys Olofsson H (2008) A circumstellar journey – physics
456:1001–1012 and chemistry of AGB stellar winds. Phys Scr
Cherchneff I (2012) The inner wind of IRC +10216 133:014028
revisited: new exotic chemistry and diagnostic for dust Petrie S (1996) On the formation of metal cyanides and
condensation in carbon stars. Astron Astrophys 545:A12 related compounds in the circumstellar envelope of
Decin L, Agúndez M, Barlow MJ et al (2010) Warm water IRC +10216. MNRAS 282:807–819
vapour in the sooty outflow from a luminous carbon Petuchowski SJ, Dwek E, Allen JE Jr, Nuth JA III
star. Nature 467:64–67 (1989) CO formation in the metal-rich ejecta of SN
Duari D, Hatchell J (2000) HCN in the inner envelope of 1987A. Astrophys J 342:406–415
cCygni. Astron Astrophys 358:L25–L28 Quintana-Lacaci G, Bujarrabal V, Castro-Carrizo A,
Gail H-P, Sedlmayr E (1988) Dust formation in stellar Alcolea J (2007) The chemical composition of the
winds. IV – heteromolecular carbon grain formation circumstellar envelopes around yellow hypergiant
and growth. Astron Astrophys 206:153–168 stars. Astron Astrophys 471:551–560
Gehrz RD (1989) Sources of stardust in the Galaxy. IAUS Schöier FL, Ramstedt S, Olofsson H, Lindvist M, Bieging
135:445–453 JH, Marvel KB (2013) The abundance of HCN in
Citric Acid Cycle 477
circumstellar envelopes of AGB stars of different

chemical types. Astron Astrophys 550:A78 Cistron
Tsuji T (1973) Molecular abundances in stellar atmo-
spheres II. Astron Astrophys 23:411–431
Woosley SE, Weaver TA (1986) The physics of Super- ▶ Gene
nova explosions. Ann Rev Astron Astrophys
24:205–256
Ziurys LM, Milam SN, Apponi AJ, Woolf NJ (2007)
Chemical complexity in the winds of the oxygen-rich
C
supergiant star VY Canis Majoris. Nature
447:1094–1097 Citric Acid Cycle
Juli Peretó
Circumstellar Grains Spain
▶ Star Dust
Keywords
Anaplerosis; Cataplerosis; Catalytic cycle; Oxi-

dative metabolism
Cirrus Cloud
William M. Irvine
Synonyms
Krebs cycle; Tricarboxylic acid (TCA) cycle
Definition
Definition
In astronomy, cirrus clouds are sources of infra-
red (IR) emission detected by the ▶ InfraRed
Citric acid cycle is a metabolic pathway often
Astronomy Satellite (IRAS), a joint project of
regarded as the final step for the complete oxida-
the USA, the UK, and the Netherlands which
tion of fuel molecules. Stoichiometrically, a 2-C
surveyed the sky at wavelengths of 12, 25,
molecule (acetyl CoA) condenses with a 4-C
60, and 100 mm. The spatial distribution of this
molecule (oxaloacetate) to yield citrate
IR emission is similar in appearance to terrestrial
(Fig. 1a). Two consecutive oxidative decarboxyl-
cirrus clouds, although the latter have nothing to
ations transform the initial 2-C unit into two CO2
do with the interstellar clouds. The astronomical
molecules. The regeneration of oxaloacetate
emission arises from interstellar dust in our gal-
closes the cycle through an oxidative process. In
axy, particularly that associated with the diffuse
addition to the electrons taken up by redox coen-
portions of interstellar clouds.
zymes (NAD(P)+ and FAD), a part of the energy
is conserved in a substrate-level phosphorylation
step yielding GTP. The citric acid cycle can also
See Also be regarded as a source of biosynthetic precur-
sors. In some organisms, the cycle operates in
▶ Infrared Astronomical Satellite reverse, reductively, thus functioning as an auto-
▶ Interstellar Dust trophic pathway (Fig. 1b).
478 Citric Acid Cycle
a Oxidative cycle b Reductive cycle

Asp, Asn
OAA OAA
2[H] 2[H]
Phosphoenolpyruvate
Malate Malate
Glucose
+ Acetyl CoA Fumarate Fumarate

2[H] 2[H]
Glyoxylate
Succinate Succinate
NTP NTP
Lys, Heme Succinyl CoA Succinyl CoA

2[H], CO2
2[H], CO2
Glu, Gln 2-Oxoglutarate 2-Oxoglutarate
2[H], CO2
2[H], CO2
Isocitrate Isocitrate
Acetyl CoA Citrate CO2 Citrate
2[H], CO2 2[H], CO2

Acetyl CoA Pyruvate Acetate Acetyl CoA Pyruvate
Citric Acid Cycle, Fig. 1 The citric acid cycle. (a) Oxi- from cycle intermediates (in gray). The glyoxylate shunt
dative cycle. An anaplerotic reaction is shown (broken is also shown (broken gray line). (b) Reductive cycle
black line), as well as some biosynthetic branches starting
History (or traffic circle) than a carrousel: There is a

permanent flux of metabolites in and out the
In an elegant series of experiments on substrate cycle. Some metabolites are used for biosynthetic
oxidation in respiratory animal tissues, purposes (e.g., 2-oxoglutarate as glutamate pre-
performed among others by Albert Szent-Gy- cursor or succinyl CoA for the biosynthesis of
örgyi (1893–1986), Carl Martius (1906–1993), heme group or lysine, see Fig. 1a). The catalytic
and Franz Knoop (1875–1946), fragments of a nature of the citric acid cycle, as well as the
sequence of the metabolic transformations were consumption of intermediates as biosynthetic
established. Those observations were completed precursors, imposes the necessity of the net syn-
by Hans A. Krebs (1900–1981) who proposed, in thesis and replacement of those intermediates.
a paper coauthored by William A. Johnson, the This process is termed anaplerosis. Anaplerotic
cyclic nature of the pathway in 1937 (Krebs and reactions include the synthesis of oxaloacetate by
Johnson 1937). pyruvate carboxylation and the synthesis of oxa-
loacetate or 2-oxoglutarate from aspartate or glu-
tamate, respectively, by transamination. On the
Overview other hand, the 4-C and 5-C skeletons derived
from amino acid catabolism must leave the
The citric acid cycle occupies a central position in cycle to be fully oxidized (this process has been
the metabolic network both as a catabolic, oxida- called cataplerosis (Owen et al. 2002).
tive process and as a supplier of biosynthetic In some autotrophic microorganisms, the
precursors. The cycle is more a roundabout citric acid cycle operates in a reverse, reductive
Clathrate Hydrate 479
way (Fig. 1b). This pathway (also known as the

Arnon-Buchanan cycle) allows the net synthesis Clathrate
of Acetyl CoA from CO2. Firstly described by
Daniel I. Arnon and Robert B. Buchanan in 1966 Henderson James (Jim) Cleaves II
as the autotrophic carbon fixation pathway in the Earth–Life Science Institute (ELSI), Tokyo
green sulfur bacterium Chlorobium limicola Institute of Technology, Meguro–ku, Tokyo,
(Evans et al. 1966), it is also present in some Japan C
proteobacteria and some members of the domain Institute for Advanced Study, Princeton, NJ, USA
Archaea (Berg et al. 2010). Blue Marble Space Institute of Science,
Washington, DC, USA
See Also of Technology, Atlanta, GA, USA
▶ Anabolism
▶ Carbon Dioxide Definition
▶ Catabolism
▶ Glycolysis In chemistry, a clathrate is an inclusion complex,
▶ Metabolism also known as a clathrate compound, cage or a
host-guest complex, in which a molecule, aggre-
gate of molecules or crystal lattice of molecules
(the “host”) noncovalently traps or encloses
References and Further Reading another, usually small, gas molecule (the
“guest”), rendering it incapable of escaping by
diffusion. The word is derived from the Latin
Autotrophic carbon fixation in archaea. Nat Rev word clatratus, meaning “with bars” or “a lattice.”
Microbiol 8:447–460 Especially important naturally occurring types
Evans MCW, Buchanan BB, Arnon DI (1966) A new of clathrates are ▶ clathrate hydrates, formed
feredoxin-dependent carbon reduction cycle in a pho-
tosynthetic bacterium. Proc Natl Acad Sci U S A
from water ice and various gases such as meth-
55:928–934 ane, ammonia or CO2, and zeolite minerals.
Krebs HA, Johnson WA (1937) The role of citric acid in
intermediate metabolism in animal tissues.
Enzymologia 4:148–156
Owen OE, Kalhan SC, Hanson RW (2002) The key role of
See Also
anaplerosis and cataplerosis for citric acid cycle func-
tion. J Biol Chem 277:30409–30412 ▶ Clathrate Hydrate
Clathrate Hydrate
Classification
William M. Irvine
▶ Taxonomy University of Massachusetts, Amherst, MA, USA
Definition
Clastation
Clathrate hydrates are ice-like solids in which a
▶ Weathering “host” hydrogen-bonded H2O lattice entraps a
480 Clay
nonpolar “guest” gas molecule such as CH4 or Clays are natural, soft, earthy, fine-grained
CO2. It has been speculated that clathrate materials (less than 2 mm); they are plastic when
hydrates are important components of many mixed with an appropriate amount of water and
outer solar system bodies including comets, the hard when fired. Clays are fundamentally com-
outer planets, and their icy moons. Methane posed of a silicon tetrahedral layer and an alumi-
clathrate hydrates are also widely distributed in num octahedral layer with water trapped between
terrestrial seafloor sediments where the ambient the silicate sheets (Guggenheim and Martin
temperature and pressure allows their existence. 1995).
It is thought that their breakdown during periods
of increasing temperature releases methane, a
strong greenhouse gas, which may contribute sig- History
nificantly to global warming.
Clays and clay minerals have been mined since
the Stone Age, when prehistoric man first discov-
ered their useful properties. Clay tablets were
See Also used as the first writing medium, and clays
sintered in fire were used to make the first
▶ Clathrate ceramic objects. In many ancient religions and
philosophies, mankind was originally created
from clay; for example, in the Bible, the first
man, Adam, whose name means “of the
▶ earth,” was made from clay.
Clay
Alicia Negrónk-Mendoza Overview

Instituto de Ciencias Nucleares, Universidad
Nacional Autónoma de México, México, Structure
Coyoacán, DF, Mexico The simple crystal structure of clay consists of
two basic components: a silicon tetrahedron-
oxygen layer (the tetrahedral silicon can be
Keywords partially replaced with Al3+ or Fe3+) and an
octahedron layer, in which an atom of alumi-
Chemical evolution; Clay; Clay minerals; Mont- num, magnesium, and/or iron is surrounded by
morillonite; Phyllosilicates six anions (oxygen or hydroxyls). The individ-
ual units are stacked in parallel plates, one above
the other, and depending upon the arrangement,
Synonyms different types of clay are produced. They are
classified first into “layer types,” according to
Argillaceous earth; Clay minerals the number of tetrahedral and octahedral sheets,
and then into “groups,” which are differentiated
by the type of isomorphic cation substitutions
Definition present. The layer types are illustrated in Fig. 1.
The 1:1 (or TO) layer type consists of one tetra-
Clay is a generic term for a mineral group of hedral sheet fused to an octahedral sheet and
complex hydrated alumino-phyllosilicates that is represented by the ▶ kaolinite group. Type
mainly form as a result of feldspar weathering 2:1 clay (or TOT) is made up of an octahedral
and low-temperature hydrothermal alteration of sheet sandwiched between two tetrahedral
many rocks. sheets; examples include kaolinite, illite, and
Clay 481
Clay, Fig. 1 Types of

clays (see text)
Tetrahedral Octahedral H2O
Kaolinite
Halloysite
C
(+)
+ + + + + +2 + +2 + + +
+
+ +
H2O
(+)
lllite
Vermiculite Chlorite
Smectite
smectites, such as ▶ montmorillonite. This com- Distribution

posite unit is continuous in two directions of a Clays are ubiquitous minerals on Earth; their
plane in the c-direction (see Fig. 2) and forms presence can likely be traced to the early stages
packets of 2–15 elementary units (Meunier of the planet’s formation. While various types of
2010). Figure 2 depicts a TOT layer-type crystal clays most likely formed on the continents of
structure. Earth in the Archean, the largest area of clay
At ▶ pH values above 2, clay particles carry a production would have been the ocean floor.
net negative charge in the space between the The genesis of continental smectites can be
silica layers, referred to as the interlayer or traced back 3 or 4 billion years ago (Odin 1988).
interlamellar channel. The distance between The presence of clays in extraterrestrial envi-
these layers varies depending upon the layers of ronments, such as ▶ meteorites, ▶ Mars, and the
water or intercalated organics (Swartzen-Allen ▶ comet Tempel 1 (Lisse et al. 2006), helps us to
and Matijevic 1974). The negative charge can understand the aqueous conditions that prevailed
originate from several different factors, including in the early stages of the history of the solar
lattice imperfections, isomorphic substitution, system, as the primary requirement for the for-
broken bonds, and exposed structural hydroxyl mation of clay minerals is the presence of liquid
groups. The presence of a positive counterion, water.
such as Na+, K+, or Ca2+, compensates for the
negative charge. The edges of the crystal are Importance
positively charged in at neutral and acidic pH Clays are important minerals in nature and in
(Swartzen-Allen and Matijevic 1974). These industry. They play an integral part of terrestrial
structural characteristics are responsible for the ▶ biogeochemical cycles. These processes influ-
special properties of clays. For example, mont- ence microbial ▶ life and the cycling of elements
morillonite can take up various types of organic on planetary surfaces. Clay also plays a role in the
molecules. The molecule may be adsorbed on the buffering capacity of the oceans.
clay lattice by cationic interchange, ion-dipole Clays are among the most important minerals
forces, ▶ van der Waals forces, or hydrogen used in the manufacturing and environmental
bonds. industries; for example, the construction of
482 Clay
Clay, Fig. 2 TOT layer-

type crystal structure
C axis
Interlayer channel (exchangable cations)
oxygen
silicon, occasionally AI
aluminon, iron,
magnesium
hydroxy
ceramics, including tiles, bricks, and pottery. In were the first functional templates (Cairns-
addition to sand and silt, clay is one of the three Smith 1966; Cairns-Smith (1982), Cairns-Smith
principal types of sediment and primarily results and Hartman 1988). Mud beds, shores, and
from the weathering of rocks and soil on the hydrothermal vents at the bottom of the sea are
surface of the Earth. all possible prebiotic scenarios in nature. In addi-
tion, these regions can be alternately wet and dry,
Importance in Chemical Evolution as in tidal estuaries. These environments stress
Many authors have proposed that clays played a the significance of multiphase systems containing
central role in ▶ chemical evolution and ▶ origin clay minerals in prebiotic processes.
of life (Negrón-Mendoza et al. 2010; Hashizume An important step from chemical to biological
2012). This role is a result of the capacity of clay evolution is the formation of cell membranes.
to adsorb organic molecules and catalyze reac- Szostak and his group (2003) reported that mont-
tions (Theng 1974; Laszlo 1987; Yariv and Cross morillonite accelerates the spontaneous conver-
2002). Clay minerals are important for several sion of fatty acid micelles into vesicles, which
purposes: (1) to concentrate the organic mole- could serve as a pathway for the prebiotic encap-
cules present in a dilute ocean by adsorption on sulation of catalytically active surfaces within
clay deposits, (2) to catalyze the polymerization membrane vesicles.
of adsorbed organic compounds, (3) to protect
these organic molecules from destruction by
ultraviolet light or other energy sources, and See Also
(4) as a probable mechanism to induce chirality
in contemporary molecules. Investigations of the ▶ Biogeochemical Cycles
role of clays in prebiotic organic synthesis range ▶ Buffer
from their capability to act as adsorbents and/or ▶ Chemical Evolution
catalysts to the controversial claim that clays ▶ Comet Tempel 1
Clean Room 483
▶ Earth Negrón-Mendoza A, Ramos-Bernal S, Mosqueira FG

▶ Earth, Formation and Early Evolution (2010) The role of clay interactions in chemical evo-
lution. In: Basiuk V (ed) Astrobiology: emergence,
▶ Environment search and detection of life. American Scientific Pub-
▶ Hydrogen Bond lishers, Los Angeles, pp 214–233
▶ Hydrothermal Reaction Odin GS (1988) The origin of clays on Earth. In: Cairns-
▶ Kaolinite Smith AG, Hartman H (eds) Clay minerals and the
origin of life. Cambridge University Press, Cambridge,
▶ Life pp 81–89
C
▶ Mars Swartzen-Allen SL, Matijevic E (1974) Surface and col-
▶ Meteorites loid chemistry of clays. Chem Rev 74:385–399
▶ Mineral Theng BKG (1974) The chemistry of clay-organic reac-
tions. Adam Hilger/Wiley, London/New York
▶ Montmorillonite Yariv S, Cross H (2002) Organo-clay complexes and
▶ Organic Molecule interactions. Marcel Dekker, New York
▶ Origin of Life
▶ pH
▶ Silicate Minerals
▶ Van der Waals Forces
▶ Water Clay Minerals
▶ Weathering
▶ Clay
▶ Phyllosilicates, Extraterrestrial
Cairns-Smith AG (1966) The origin of life and the nature

of the primitive gene. Theor Biol 10(1):53–88
Cairns-Smith AG (1982) Genetic takeover – and the min-
eral origins of life. Cambridge University Press,
Clean Room
Cambridge
Cairns-Smith AG, Hartman H (1988) Clay minerals and Gerda Horneck
the origin of life. Cambridge University Press, DLR German Aerospace Center, Institute of
Cambridge
Aerospace Medicine, Radiation Biology,
Guggenheim S, Martin RT (1995) Definition of clay and
clay minerals: join report of the AIPEA nomenclature Köln, Germany
and CMS nomenclature committees. Clay Clay Miner.
43:255–256
Hanczyc MM, Fujikawa SM, Szostak, JW (2003) Exper-
imental models of primitive cellular compartments:
Definition
encapsulation, growth, and division. Science 302
(5645):618–22 A clean room is a workplace, generally a room,
Hashizume H (2012) Role of clay minerals in chemical where the air is permanently filtered to remove
evolution and the origins of life. In: Valaškova M,
the particles. In some cases, the temperature and
Simha Martynkova G (eds) Clay minerals in
nature – their characterization, modification and appli- atmospheric humidity are also controlled while
cation. ISBN 978-953-51-0738-5. InTech, Rijeka, the number of particles is monitored. The access
Croatia, Open Access, pp 192–208 is strictly limited to trained personnel entering
Laszlo P (1987) Chemical reactions on clays. Science
through a double door after they dress with pro-
235:1473–1477
Lisse CM, VanCleve J, Adams AC, Ahearn MF, tective garment to avoid the spreading of skin
Fernández YR, Farnham TL, Armus CML, Grillmair particles and hair. In such clean rooms, the labo-
CJ, Ingalls J, Belton MJS, Groussin O, McFadden LA, ratory hardware, the lab wear of the personnel,
Meech KJ, Schultz PH, Clark BC, Feaga LM, Sunshine
and the number of people working simulta-
JM (2006) Spitzer spectral observations of the deep
impact ejecta. Science 313:635–640 neously are also specified to limit the spreading
Meunier A (2010) Clays. Springer, Berlin of dust and particles. Clean rooms are used in
484 Cleanliness
industry to perform the tasks requiring the max- be considered. The chemical cleanliness refers to
imum cleanliness such as the production of maximal specified amounts of each chemical per
microprocessors, hard disk drives, and precision unit of surface or volume.
optical elements.
Minimizing the number of particles minimizes
also the number of airborne ▶ microorganisms See Also
and subsequently the contamination of the
surfaces. ▶ Bioburden
Clean rooms are required for the assembly of ▶ Planetary Protection
the satellites and of the interplanetary probes ▶ Sterilization
following planetary protection policy. In prac-
tice, the clean rooms are often known by the
number of particles with size under 0.5 mm by
cubic feet (class 10,000, class 1,000, class
Cleverness
100, etc.). A strict definition is proposed by the
International Organization for Standardization
▶ Intelligence, Evolution of
(ISO) in the standard ISO 14644-1. This defini-
▶ Intelligence
tion applies also to small working area limited to
a tent or to working cabinets.
Cloning
See Also
Juli Peretó
▶ ISO
▶ Microorganism
Spain
Definition
Cleanliness
Cloning is the process of obtaining replicative
Catharine A. Conley molecules, cells, or organisms that are geneti-
NASA Headquarters, Washington, DC, USA cally identical to a common ancestor. DNA
recombinant techniques – or ▶ genetic
engineering – allow the synthesis, replication,
Definition and expression of DNA fragments of any size,
up to complete synthetic molecules the size of a
For ▶ planetary protection, the cleanliness of an small bacterial genome, in appropriate receptor
environment or spacecraft hardware is the level cells. The cloning of cells and organisms – e.g.,
of contamination by something which is not plants and animals – using the methods of tissue
wanted for the achievement of a goal. The max- culture and in vitro cell differentiation is also
imum amount of the unwanted item is generally possible.
specified. For instance, the biological cleanliness
can be measured by the number of remaining
microorganisms after the cleaning process. See Also
Depending on the final goal, the living microor-
ganisms, the dead microorganisms, or both could ▶ Amplification (Genetics)
Clouds 485
first process needs a highly supersatured vapor

Clouds phase, while the latter one already operates at
very low supersaturations but requires the a priori
Mark S. Marley1, Lisa Kaltenegger2 and Daniel presence of some form of seed particles (e.g.,
Kitzmann3 ▶ aerosol particles). The resulting initial distri-
1
NASA Ames Research Center, Moffett Field, bution of nuclei undergoes numerous microphys-
CA, USA ical processes, such as growth, evaporation, C
2
Cornell University, Ithaca, NY, USA coagulation, or sedimentation, which determine
3
University of Bern, Bern, Switzerland the temporal and spatial evolution of the cloud’s
particle size distribution (Pruppacher and Klett
1997). Because exoplanetary atmospheres can
Keywords span a very wide range of chemical compositions
as well as temperature and pressure conditions, a
Albedo; Atmosphere; Clouds; Extrasolar planets; large number of species may form clouds.
Spectra; Spectroscopy Jupiter’s atmosphere provides a point of
departure for understanding the diversity of
giant planet atmospheres that we expect to
Synonyms encounter outside of the solar system. Assuming
we start at a temperature of about 2,000 K
Condensate layer (Fig. 1), the first constituents to condense in a
rising gas parcel are refractory oxides such as
perovskite and corundum, followed by various
Definition magnesium silicates including enstatite and
forsterite. As we move upward in the atmosphere,
Clouds are condensates (either liquid or solid) of the temperature continues to fall and eventually
gaseous atmospheric constituents, which play an water clouds form, removing H2O from the gas
important role in planetary atmospheres. Since phase. Above the water clouds, the atmosphere
clouds scatter and absorb incident stellar radia- continues to cool until ammonia clouds form. It is
tion as well as emergent thermal radiation and the ammonia clouds of Jupiter, dusted by various
release ▶ latent heat during their formation, they photochemical pollutants, that we see reflecting
control both the appearance and thermal structure sunlight back from the planet. In a warmer Jupi-
of a planet. Furthermore, a cloud deck can both ter, the atmosphere would never become satu-
limit the depth an external observer can “see” into rated in water and ammonia vapor, and these
an atmosphere and thus hide molecular species, species would stay in the gas phase, thus remov-
and it can alter a planet’s ▶ albedo. For such ing the bright clouds and substantially altering
reasons, a basic understanding of the role clouds the appearance and color of the planet.
play is required to interpret exoplanet spectra or Direct evidence for the presence of clouds has
to obtain consistent temperature profiles and sur- been found in transmission spectra of several
face temperatures. transiting extrasolar gas giants. Currently, the
best example for the occurrence of high-altitude
clouds is the transiting hot Jupiter HD 189733b
Overview (Sing et al. 2011). The possible existence of clouds
is also debated for super-Earth planets, such as GJ
Clouds can form when the partial pressure of a 1214b (Morley et al. 2013), although better obser-
chemical species exceeds its saturation vapor vational constraints are required to obtain more
pressure. The initial distribution of cloud nuclei accurate estimates on the clouds’ properties.
is formed by either homogenous or heteroge- Clouds play also a crucial role for terrestrial
neous nucleation from the gas phase, where the planets. For habitable planets, the most important
486 Clouds
▶ habitable zone, as they can both raise the Bond

albedo, thus cooling the surface (Kasting 1991)
NH3 and limit thermal emission, exhibiting a ▶ green-
NH4HS house effect (Kitzmann et al. 2010; Forget and
Pierrehumbert 1997).
H2O
CH4 gas Future Directions
CsCl Clouds are intrinsically difficult to model from a

RbCl priori physical considerations (see, e.g.,
Ackerman and Marley 2001; Helling
KCl et al. 2008; Marley et al. 2013). The detailed
behavior of the terrestrial cloud cover as a func-
tion of atmospheric temperature is the leading
source of uncertainty in global climate models.
LiF
Li2S Predicting cloud behavior for extrasolar planets,
Na2S including such issues as particle sizes, vertical
distribution, and any horizontal patchiness is,
CO gas therefore, difficult. Accounting for the effects of
these clouds has proven challenging, and such
Mg-silicates
issues must be carefully considered when spa-
iron metal liquid tially resolved exoplanet spectra eventually
become available.
CO gas
perovskite corundum
See Also
Clouds, Fig. 1 Cloud structure expected on a Jupiter-like
planet depending on its temperature (hot: bottom, cool: ▶ Adiabatic Processes
top) (Figure modified from Lodders (2004)) ▶ Albedo
cloud species are water droplet and carbon diox- ▶ Exoplanets, Modeling Giant Planets’
ide and water ice clouds. Typically, the reflectiv- Atmospheres
ity of water clouds in the visible and near-infrared ▶ Greenhouse Effect
wavelength range is high in comparison to sur- ▶ Habitable Planet, Characterization
face features. Thus, a cloudy planet is brighter ▶ Habitable Zone
and has a higher albedo than either a rocky, ▶ Jupiter
airless world or a planet with a deep, clear atmo-
sphere. Cold, high-level clouds can limit the
height from which thermal flux is emitted in an References and Further Reading
atmosphere and thus hide underlying regions
Ackerman A, Marley M (2001) Precipitating condensa-
which might otherwise produce significant spec-
tion clouds in substellar atmospheres. J Astrophys
tral features. They can also directly influence 556:872–884
important spectral features of molecules (e.g., Forget F, Pierrehumbert RT (1997) Warming early Mars
the potential ▶ biomarker ozone) originating with carbon dioxide clouds that scatter infrared radia-
tion. Science 278(5341):1273
from above the cloud layer (Vasquez
Helling C et al (2008) Comparative study of dust cloud
et al. 2013). Clouds must be considered in any modelling for substellar atmospheres. Mon Not
calculation of the location and extension of the R Astron Soc 391(4):1854–1873
CNES 487
Kaltenegger L, Jucks K, Traub W (2007) Spectral evolu- Definition

tion of an Earth-like planet. Astrophys J 658:598
Kasting J (1991) CO2 condensation and the climate of
early Mars. Icarus 94:1–13, ISSN 0019-1035 Radio astronomers have detected the cyanide
Kitzmann D et al (2010) Clouds in the atmospheres of ▶ anion, CN, in the circumstellar envelope of
extrasolar planets. I. Climatic effects of multi-layered the evolved carbon-rich star CW Leo (also called
clouds for Earth-like planets and implications for hab- IRC+10216). It is the smallest molecular anion
itable zones. Astron Astrophys 511:A66
Lodders K (2004) Brown Dwarfs–Faint at Heart, Rich in (2 atoms) thus far identified in such regions C
Chemistry. Science 303:323 (the anions C3N and C5N, for example, had
Marley MS et al (2013) Clouds and hazes in exoplanet previously been observed). For references, see
atmospheres. In: Mackwell S, Bullock M, Harder ▶ Molecules in Space.
J (eds) Comparative climatology of terrestrial planets.
University of Arizona Press, Tucson
Morley C et al (2013) Quantitatively assessing the role of
clouds in the transmission spectrum of GJ 1214B. See Also
Astrophys J 775:A33
Pruppacher H, Klett J (1997) Microphysics of clouds and
precipitation, 2nd edn. Kluwer, Dordrecht ▶ Circumstellar Chemistry
Sing DK et al (2011) Hubble space telescope transmission ▶ Molecules in Space
spectroscopy of the exoplanet HD 189733b: high- ▶ Radio Astronomy
altitude atmospheric haze in the optical and near-
ultraviolet with STIS. Mon Not R Astron Soc
416:1443
Vasquez M et al (2013) Infrared radiative transfer in
atmospheres of Earth-like planets around F, G, K,
and M-type stars – II. Thermal emission spectra CNES
influenced by clouds. Astron Astrophys 557:A46
Michel Viso
CNES/DSP/SME, Vétérinaire/DVM,
CMB
▶ Cosmic Background Radiation Synonyms
French Space Agency
CN
Definition
▶ Cyanogen Radical
The French Space Agency was established in
1961 and was tasked to implement and coordinate
space policy as decided by government, through
its own teams and with national and international
CN- partners in science and industry. France is a
founding member of the European Space Agency
William M. Irvine (ESA) and the Centre National d’Études
University of Massachusetts, Amherst, MA, USA Spatiales (CNES) represents the country in its
council and boards. The 2,400 employees work-
ing in 2014 at CNES are spread among the four
Synonyms main centers. The headquarters are located down-
town Paris, as well as the directorate for
Cyanide anion; Cyanide ion launchers now co-located with the ESA launcher
488 CNO Cycle
directorate. The large technical center is located There are three such cycles, each converting four
in Toulouse in the south of France. Finally, CNES protons to one 4He nucleus and releasing about
is also in charge of the launch base of Kourou 6.6 MeV/nucleon or 5 1018 erg/g. At temperatures
located in French Guyana. This base is the Euro- higher than 17 106 K, encountered in stars more
pean spaceport from which the European massive than 1.3–1.5 MJ, depending on
launchers are operated since 1973. Besides metallicity, rotation, etc., the CNO cycle is the
Ariane V, Vega rockets and Russian-built Soyuz dominant source of energy production, while in
rockets are launched also from there. The CNES lower-mass stars, the proton-proton ▶ (p-p) chains
is supporting through grants and industrial con- dominate. Although the sum of C + N + O abun-
tracts most of the French activities related to dances is conserved, 12C and 16O are converted to
14
space prototyping either commercial or institu- N, thus 14N is the main product of the CNO cycle
tional applications, supporting science in space or from the chemical evolution point of view.
activities related to defense and security.
CNES was involved in manned space flight
through cooperation with the Soviet Union lead- History
ing to the flight of Jean-Lou Chretien in 1981 and
with the United States leading to the flight of The CNO cycle is also called the Bethe-Weizs-
Patrick Baudry in 1985. Since then, several äcker cycle. Hans Bethe won the 1967 Nobel
flights and experiments related to exobiology Prize in physics for his 1938 discovery of energy
flew onboard the Mir space station (▶ COMET, production in stars.
Exobio) as well as the US space shuttle. Since the
international space station era, CNES is
supporting manned space flight activities mainly
under the auspices of ESA. See Also
CNES is also playing a major role in the
exploration of the solar system through numerous ▶ High-Mass Star
contributions to missions sponsored by the ▶ P-P Chains
United States, the Soviet Union then Russia,
India, Japan, and China. France is the first con-
tributor to ESA mixing the mandatory and the
optional programs. CNSA
Florence Raulin-Cerceau
Maı̂tre de Conférences, Centre Alexandre Koyré
(UMR 8560-CNRS/EHESS/MNHN/CSI)
CNO Cycle Muséum National d’Histoire Naturelle, Brunoy,
France
Nikos Prantzos
Definition
Definition The China National Space Administration

(CNSA) was established in 1993, as a govern-
The CNO cycle is a series of nuclear reactions mental institution to develop and fulfill China’s
converting H to He and using C, N, and due international obligations, with the approval
O isotopes as catalysts; thus, the total amount of by the Eighth National People’s Congress of
these latter species is not affected by the operation. China (NPC). Then CNSA was assigned as an
CO2 Ice Clouds (Mars) 489
internal structure of the Commission for Science,

Technology and Industry for National Defense CO2 Ice Cap (Mars)
(COSTIND).
The China National Space Administration François Forget
assumes the main responsibilities for signing Institut Pierre Simon Laplace, Laboratoire de
governmental agreements in the space area on Météorologie Dynamique, UMR 8539,
behalf of organizations; intergovernmental scien- Université Paris 6, Paris, France C
tific and technical exchanges; and also being in
charge of the enforcement of national space pol-
icies and managing the national space science, Definition
technology, and industry.
CNSA is backed by numerous independent CO2 ice caps on ▶ Mars are mantles of CO2 ice
organizations, ruling the industry for launchers deposited above 50 latitude in both hemispheres
and satellites, manned space flights, etc. The during fall and winter. They form by condensa-
China Aerospace Corporation, the three launch tion of CO2 gas, the main constituent of the atmo-
bases, the China Astronaut Research and Train- sphere, and can reach thicknesses of 50 cm to
ing Center, and the Beijing Aerospace Command 1 m. Their surface temperature is controlled by
and Control Center are some of these entities. solid-gas equilibrium with the atmosphere and
China while developing the manned space ranges between 142 and 150 K. In the northern
flights is developing science exploration of the hemisphere, the CO2 ice cap completely disap-
▶ Moon (Chang’e 1 and 2) and ▶ Mars pears during spring. In the southern hemisphere,
(▶ Yinghuo-1). the CO2 ice does not completely sublime, leaving
Up to now, China has signed governmental a perennial ▶ polar caps (Mars) 300 km across
space cooperation agreements with Brazil, and several meters thick near the south pole.
Chile, France, Germany, India, Italy, Pakistan,
Russia, Ukraine, the United Kingdom, the United
States, and some other countries. See Also
▶ Carbon Dioxide
▶ Mars
See Also ▶ Polar Caps (Mars)
▶ Yinghuo-1
CO2 Ice Clouds (Mars)
François Forget
CO Institut Pierre Simon Laplace, Laboratoire de
Météorologie Dynamique, UMR 8539,
▶ Carbon Monoxide Université Paris 6, Paris, France
Definition
CO2 Clouds composed of CO2 ice form when CO2 gas

(the main constituent of the ▶ Mars atmosphere)
▶ Carbon Dioxide condensates in the atmosphere. CO2 ice clouds
490 Coagulation in Planetary Disks
have been observed in the lower Martian atmo- coagulation refers to a statistical model for parti-
sphere during the northern and southern winter cle growth in ▶ protoplanetary disks.
polar nights (Pettengill and Ford 2000) and in
Mars’s mesosphere (80–100 km altitude) near
See Also
the equator (Montmessin et al. 2007). Thick
CO2 ice clouds may have been present in the
▶ Planetesimals
▶ planet’s denser early atmosphere, more than
▶ Planet Formation
3.5 billion years ago. They could have contrib-
▶ Protoplanetary Disk
uted to warming the surface through the process
of the scattering ▶ greenhouse effect (Forget and
Pierrehumbert 1997); however, the extent of this
effect is debated.
Coagulation, Interstellar Dust Grains
See Also William M. Irvine

▶ Carbon Dioxide
▶ Mars Definition
▶ Planet
Low-energy collisions of interstellar grains can
result in the growth of the grains or grain aggre-
gates, a process sometimes referred to as coagu-
Forget F, Pierrehumbert RT (1997) Warming early Mars lation. The structure of the grains or aggregates
with carbon dioxide clouds that scatter infrared radia- presumably depends on the size and structure of
tion. Science 278:1273–1276 the colliding particles and possibly on the pres-
Montmessin F, Gondet B, Bibring JP, Langevin Y, ence or absence of an icy grain mantle.
Drossart P, Forget F, Fouchet T (2007) Hyper-spectral
imaging of equatorial CO2 ice clouds on Mars by
OMEGA on Mars Express. J Geophys Res 112:E11,
CiteID E11S90 See Also
Pettengill GH, Ford PG (2000) Winter clouds over the
north Martian polar cap. Geophys Res Lett
27:609–613 ▶ Interstellar Dust
Coagulation in Planetary Disks

Code
Rory Barnes
Astronomy Department, University of Hugues Bersini
Washington, Seattle, WA, USA IRIDIA, Université Libre de Bruxelles, Brussels,
Belgium
Definition
Keywords
Coagulation is the first stage in ▶ planet forma-
tion in which small solid grains collide and grow. ASCII code; Genetic code; Programs; Sema-
In the context of planet formation models, phore; Turing machine
Code 491
Definition from one zone of the computer to another). The

ambiguity recognized by many biologists in the
At first and most basically, a code defines a map- meaning of “▶ gene” can be considered in the
ping between one type of information and light of these basic and extended meanings. As a
another. For instance, the ASCII code in com- matter of fact, a gene can be seen not only as
puter science is the binary translation of our coding for the associated protein but, more and
alphabet, mapping, for instance, the lowercase more in a richer manner, as indexing a whole and C
character “a” onto “1100001” and making possi- sophisticated biological process that might even
ble for computers to store, treat, and exploit infor- execute differently according to the surrounding
mation expressed in words. “Code” is one of environment (rendering the distinction between
those terms that testify to the strong intellectual innate and acquired characteristics much more
connection and mutual conceptual enrichment subtle and interesting). When Richard Dawkins
that, since Turing, Von Neumann, and the former evolves his biomorphs (Dawkins 1996 [1986]), as
systemic and cybernetic schools, has always described in his book “The Blind Watchmaker,”
existed between biology and computer science. the sophistication of the obtained creatures is not
Interestingly enough, the different meanings of so much a reflection of the genetic code used by
this term in computer science is mirrored by a him but rather appears as an outcome of the
progressive semantic enrichment also of its use in recursive program simply parameterized by this
biology. code. The genes boil down to a simple parame-
terization of a very sophisticated process respon-
sible for the major part of the resulting
Overview complexity. A mapping is still in place, but the
action/process/object mapped by the gene (such
The most celebrated biological code is without as the action/process/object mapped by a soft-
doubt the “genetic” one which maps triplets of ware instruction in a program) is far from
four possible ▶ nucleotides (“A,” “T,” “G,” “C”) static. It turns out to be a very rich one-to-many
onto 1 of the 20 ▶ amino acids found in the mapping, from one gene to many potential
▶ proteins of living organisms. For instance, the dynamical processes, the latter being capable to
triplet “AAT” is mapped onto the “Leucine” behave in a very sophisticated manner and very
amino acid. Although the ▶ genetic code is not sensitively to the surrounding environment. In
as arbitrary as the ASCII one, still a mapping the most complex cases, these dynamic processes
table is really what defines the coding in both can even interfere back on the genes, requiring
cases. then to include the temporal dimension in the
There is an extended, more sophisticated def- definition of this mapping.
inition of “code” in computer science, such as
when a programmer is writing a “code” to be
executed by his computer. The mapping is not
anymore between one type of representation and See Also
another, but pieces of the code serve as an index
to executable processes, such as when a traffic ▶ Artificial Life
light turns red requiring the driver to stop his car. ▶ Genetic Code
An essential part of elementary instructions in
computers is indeed expressed according to a
specific code (like “add” or “load”), making
both the central process unit and the memory
participate in the execution of a whole process Dawkins R (1996 [1986]) The blind watchmaker. WW
(such as adding numbers or copying information Norton, New York. ISBN 0-393-31570-3
492 Codon
Definition
Codon
Coenzyme is any organic, low-molecular-mass,
Juli Peretó freely dissociable factor, which is essential for
Institut Cavanilles de Biodiversitat i Biologia the activity of an enzyme. For example, some
Evolutiva, Universitat de València, València, Spain dehydrogenases require NAD+ as electron carrier
and some reductases use NADPH.
Definition See Also
A codon is each of the nonoverlapping nucleotide ▶ NADH, NADPH

triplets in the coding sequence of an mRNA,
which specifies an amino acid, following the
equivalences of the ▶ genetic code.
Coevolution
See Also Stéphane Tirard

Centre François Viète d’Histoire des Sciences et
▶ Anticodon des Techniques EA 1161, Faculté des Sciences et
▶ Genetic Code des Techniques de Nantes, Nantes, France
▶ Ribosome
▶ RNA
History
▶ Translation
The concept of coevolution was developed in
1964 by Paul R. Ehrlich (1932) and Peter
H. Raven (1936). It consists in the evolutionary
interaction between two species. Coevolution
Codon Table
notably concerns the linked evolution in the
some host-parasite relationship, each of them
▶ Genetic Code
interacts with the evolution of the other, and the
relationship stays stable.
In 1973, Leigh Van Valen (1935–2010) pro-
Co-elution
posed the famous parabola of the “Red Queen’s
model,” inspired by Lewis Carroll’s novel,
▶ Chromatographic Coelution
Through the Looking-Glass, and What Alice
Found There (1871). Alice and the Red Queen
run together and they stay under the same tree all
Coenzyme
the time. The Red Queen explains to Alice:
“Now, here, you see, it takes all the running you
Juli Peretó
can do, to keep in the same place.” This sentence
resumes the idea of coevolution in which the two
species have evolved simultaneously to maintain
Spain
their relationship in the same state.
Synonyms See Also
Cofactor ▶ Evolution, Biological

Collection en Orbite de Matériel Extra Terrestre 493
This formula can be written as

Cofactor Pc ¼ N 0 RPS PI PR Pg .
In ▶ planetary protection, the formulation is
Juli Peretó often used by determining the initial number of
Institut Cavanilles de Biodiversitat i Biologia microorganisms (N0) that could be present on or
Evolutiva, Universitat de València, València, Spain in a spacecraft and multiplying this number by an
appropriately selected set of factors representing C
the probability this number could be reduced. The
Definition first reduction in proportion (R) is depending on
various parameters, including conditions to
Cofactor is a nonprotein (organic or inorganic) which the spacecraft is exposed both before and
factor necessary for the activity of an ▶ enzyme. after launch. Then, while present on the space-
This factor can be firmly bound to the enzyme craft, the microorganisms have to reach the sur-
(i.e., prosthetic group, e.g., ▶ cytochromes) or face of the planet. The value PI describes the
freely dissociable (i.e., ▶ coenzyme, e.g., NAD probability of the spacecraft to hit the planet.
(P)H). This value is ranging from 105 for a satellite
up to one for a landing probe. The probability for
a microbe to be released (PR) in the environment
See Also while the spacecraft is on the ground is generally
set to one in case of crash landing. The probabil-
▶ Coenzyme ity of growth (Pg) in the particular extraterrestrial
▶ Cytochromes environment is also included in the calculation,
▶ Enzyme but for targets with liquid water is assumed to be
▶ NADH, NADPH one. The “probability of contamination” is taken
to be equivalent to the fractional number of
organisms (e.g., 1 104) that could be present
on a spacecraft after various reduction factors are
included.
Cognition
▶ Intelligence, Evolution of See Also

▶ Intelligence
▶ Bioburden
▶ Microorganism
Coleman-Sagan Equation
Catharine A. Conley
NASA Headquarters, Washington, DC, USA Sagan C, Coleman S (1965) Spacecraft sterilization stan-
dards and contamination of Mars. Astronaut Aeronaut
3(5):1–22
Definition
The Coleman-Sagan equation is used for

establishing the probability of contaminating Collection en Orbite de Matériel
another planetary body by Earth microorganisms Extra Terrestre
and was first published by S. Coleman and
C. Sagan in 1965. ▶ COMET (Experiment)
494 Colonization, Biological
including planets, by the human species. Biolog-

Colonization, Biological ical colonization is a dynamic process that begins
when unoccupied habitats, territories, or niches
Silvano Onofri become available or when organisms acquire the
Department of Ecological and Biological ability to survive and reproduce under environ-
Sciences, University of Tuscia, Viterbo, Italy mental conditions of new niches, by a process of
adaptation (Fig. 1).
Keywords
Overview
Archaea; Dormant state; Ecological niche;
Eukarya; Extremotolerance; Habitat; Microbial The colonization of new habitats involves a suc-
community; Origin of life; Panspermia cession of biological communities that can be stud-
ied and, to some extent, predicted. In many cases,
the biological colonization of new habitats and
Synonyms territories begins with microorganisms, which has
particular relevance to astrobiology. All known life
Settlement forms are based on ▶ carbon chemistry and are
related to the availability of ▶ water. Organisms
are able to colonize all environments that fall
Definition within the limits of habitability, i.e., conditions of
low (Kappen et al. 1996; Cavicchioli 2006) to high
Colonization is the occupation of a ▶ habitat or temperatures (up to 122 C, Kashefi and Lovley
territory by a biological community or of an 2003; Takai et al. 2008), low water availability
ecological niche by a single population of a spe- (Billi and Potts 2002; Leong et al. 2011), extreme
cies. Biological colonization relates to all spe- acidity (Amaral Zettler et al. 2002; Schleper et al.
cies, from microbes – including bacteria, 1996) or alkalinity, high pressure, salinity (Oren
archaea, and ▶ fungi – to more complex organ- 2000; 2002; Pikuta and Hoover 2007), and radia-
isms, like plants and animals. The term also tion (Billi et al. 2000; Bagwell et al. 2008;
applies to the occupation of new territories, Dadachova and Casadevall 2008). We find life,
Colonization, Biological,
Fig. 1 Cryptoendolithic
(hidden within rock)
microbial community
colonizing sandstone,
McMurdo Dry Valleys,
continental Antarctica (bar
1 cm) (Copyright # 2009
Laura Zucconi (permission
obtained))
Colonization, Biological 495
Fig. 2 A “lichen’s grass”
(Usnea antarctica) in King
George Island, Antarctica
(bar 10 cm) (Copyright #
2001 Silvano Onofri)
Fig. 3 Mosses colonizing
rocks in King George
Island, Antarctica (bar
Silvano Onofri)
primarily microbial, in hot springs on the ocean colonization is thought to be possible wherever
floor, within the crust to depths over 3,000 m water could be available, such as in the perma-
(Cockell 2003), in deserts (Friedmann 1982) and frost of ▶ Mars or on ▶ Europa Jupiter’s moon.
on the highest mountains, in the Arctic and Ant- The notion of biological colonization beyond
arctic ▶ permafrost (Rivkina et al. 2000), high in Earth leads to speculation about the transfer of
the stratosphere, in salt crystals, in acidic (up to pH life from one celestial body to another. Today, we
0 or negative) to very basic waters (Schleper et al. know that the conditions suitable for the origin of
1996; Takai et al. 2001), and in sites contaminated life could have existed on Mars and perhaps on
by nuclear radiation (Dadachova and Casadevall other planets. Many organisms are capable of
2008) (Figs. 2 and 3). surviving in a dormant state in conditions far
The limits of metabolically active life define more prohibitive than those suitable for active
the habitability of celestial bodies. Biological metabolism. Some organisms, e.g., bacteria
496 Colonization, Biological
Fig. 4 A higher plant
(Colobanthus quitensis)
colonizing soil in King
George Island, Antarctica
(bar 10 cm) (Copyright #
2001 Silvano Onofri)
Fig. 5 Lichens
(Umbilicaria and
Xanthoria) and higher
plants (Colobanthus
quitensis and Deschampsia
antarctica) colonizing
rocks in King George
Island, Antarctica (bar
Silvano Onofri)
(Horneck et al. 1994), microfungi (Onofri beginning the biological colonization of new ter-
et al. 2008; 2012), and lichens (Sancho ritories (Horneck and Baumstark-Khan 2002)
et al. 2007), have been shown to withstand (Figs. 4 and 5).
space vacuum, temperatures, and radiation.
Thus, they are valuable model organisms for
studying possible extraterrestrial life. Bacterial See Also
spores within rocks are able to withstand the
shock of an ▶ asteroid impact and could therefore ▶ Antarctica
have been ejected from the parent planet and ▶ Asteroid
could also have survived the landing process. ▶ BIOPAN
Life could thus have been transferred from one ▶ Black Smoker
planet to another (▶ Lithopanspermia), ▶ Carbon
Colonization, Biological 497
▶ Deep-Sea Microbiology Horneck G, B€ ucker H, Reitz G (1994) Long-term survival

▶ Deep Subsurface Microbiology of bacterial spores in space. Adv Space Res 14:41–45
Kappen L, Schroeter B, Scheidegger C, Sommerkorn M,
▶ Epilithic Hestmark G (1996) Cold resistance and metabolic
▶ Europa activity of lichens below 0 C. Adv Space Res
▶ Expose 12:119–128
▶ Extreme Environment Kashefi K, Lovley DR (2003) Extending the upper tem-
perature limit for life. Science 301:934
▶ Fungi Leong Sl-L, Pettersson OV, Rice T, Hocking AD,
C
▶ Habitable Zone Schnurer J (2011) The extreme xerophilic
▶ Habitat mould Xeromyces bisporus – growth and competition
▶ Lithopanspermia at various water activities. Int J Food Microbiol
145:57–63
▶ Mars Onofri S, de la Torre R, de Vera JP, Ott S, Zucconi L,
▶ Meteorites Selbmann L, Scalzi G, Venkateswaran K, Rabbow E,
▶ Panspermia Sánchez Inigo FJ, Horneck G (2012) Survival of rock-
▶ Permafrost colonizing organisms after 1.5 years in outer space.
Astrobiology 12:508–516
▶ Rock Onofri S, Barreca D, Selbmann L, Isola D, Rabbow E,
▶ Space Environment Horneck G, de Vera JPP, Hatton J, Zucconi L (2008)
▶ Space Vacuum Effects Resistance of Antarctic black fungi and cryptoen-
▶ Spallation Zone dolithic communities to simulated space and Mars
conditions. Stud Mycol 61:99–109
▶ Spore Oren A (2002) Molecular ecology of extremely halophilic
▶ UV Radiation, Biological Effects Archaea and Bacteria. Fems Microbiol Ecol 39:1–7
▶ Water Oren A (2000) Diversity of halophilic microorganisms:
environments, phylogeny, physiology, and applica-
tions. J Ind Microbiol Biotechnol 28:56–63
Pikuta EV, Hoover RB (2007) Microbial extremophiles at
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Chroococcidiopsis. Appl Environ Microbiol transvaalensis gen. nov., sp nov., an extremely
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Cavicchioli R (2006) Cold-adapted archaea. Nat Rev can gold mine. Int J Syst Evolut Microbiol
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Cockell C (2003) Impossible extinction. Cambridge Uni- Takai K, Nakamura K, Toki T, Tsunogai U, Miyazaki M,
versity Press, Cambridge Miyazaki J, Hirayama H, Nakagawa S, Nunoura T,
Dadachova E, Casadevall A (2008) Ionizing radiation: Horikoshi K (2008) Cell proliferation at 122 degrees
how fungi cope, adapt, and exploit with the help of C and isotopically heavy CH4 production by a hyper-
melanin. Curr Opin Microbiol 11:525–531 thermophilic methanogen under high-pressure cultiva-
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life. Springer, Berlin doi:10.1038/srep05114
498 Color Excess
longer, standard wavelength: this is a quantitative

Color Excess measure of a star’s color. A positive color index
indicates a star redder (generally cooler) than an
Daniel Rouan A0 star, such as Vega, and a negative one, a bluer
LESIA, Observatoire Paris-Site de Meudon, (hotter) star.
Meudon, France
See Also
Definition
▶ Color Excess
The color excess is the difference between the ▶ Magnitude
observed ▶ color index of a star and the intrinsic
color index predicted from its spectral type. It is a
quantity that is always positive and that gives a
measure of the absorption of starlight by the
intervening ▶ interstellar medium. The apparent Color-Magnitude Diagram
reddening of the starlight comes from a stronger
absorption in the blue than in the red by dust ▶ Hertzsprung-Russell Diagram
particles. Generally, it’s the excess of the [B-V]
color index of the Johnson photometric system
which is used: it is denoted EB-V. The interstellar
extinction, denoted Av, is directly proportional to
EB-V. (Av 3 EB-V). Column Density
Daniel Rouan
See Also LESIA, Observatoire Paris-Site de Meudon,
Meudon, France
▶ Color Index
▶ Dust Grain
▶ Interstellar Medium Definition
▶ Magnitude
▶ Reddening, Interstellar The column density between two points in a
medium is the projected number density of a
given species (H atoms, dust particles) contained
in a cylinder whose length is the distance between
the two points. The unit is m2. When applied to
Color Index the interstellar matter lying between an object
and the Earth, the hydrogen column density is
Daniel Rouan proportional to the extinction, with the approxi-
LESIA, Observatoire Paris-Site de Meudon, mate relation: Av = 5 1026 N(H), where N(H)
Meudon, France is the column density or projected number of
H atoms per square meter and Av is the visual
extinction.
Definition
A color index is the difference between the See Also

▶ magnitude of a star measured at one standard
wavelength and the magnitude at another, albeit ▶ Extinction, Interstellar or Atmospheric
Combustion 499
Definition
Combinatorial Library
A combinatorial nucleic acid library is a large
Burckhard Seelig pool of DNA or RNA molecules used as the
Department of Biochemistry, Molecular Biology starting point of an in vitro selection or evolution
and Biophysics & BioTechnology Institute, experiment. Such libraries usually contain
University of Minnesota, St. Paul, MN, USA 1012–1016 nucleic acid molecules showing a cen- C
tral region – typically, 20–100 nucleotides long –
of random or mutagenized sequence, flanked by
Definition two primer-binding sites of defined sequence.
Since 1990, combinatorial nucleic acid libraries
Combinatorial libraries are mixtures of large have been used to isolate specific nucleic acid
numbers of compounds. To study the origin ligands for a variety of target molecules –
and evolution of biomolecules such as nucleic aptamers – , as well as nucleic acid-based cata-
acids or proteins, vast combinatorial libraries lysts for different reactions – ribozymes and
have been synthesized that contain up to 1015 deoxyribozymes–.
different random RNA molecules or up to 1013
random polypeptides. These libraries are used in
conjunction with in vitro selection techniques
such as SELEX (systematic evolution of ligands See Also
by exponential enrichment) or mRNA display to
test the hypothesis of the emergence of func- ▶ Aptamer
tional biomolecules from random sequence ▶ Evolution, In Vitro
space. ▶ Evolution, Molecular
▶ Nucleic Acids
▶ Ribozyme
▶ Sequence
See Also
▶ mRNA Display
▶ Protein Combustion
▶ RNA
Japan
Combinatorial Nucleic Acid Library Institute for Advanced Study, Princeton, NJ, USA
Carlos Briones Washington, DC, USA
Centro de Astrobiologı́a (CSIC/INTA), Consejo Center for Chemical Evolution, Georgia Institute
Superior de Investigaciones Cientı́ficas, Madrid, of Technology, Atlanta, GA, USA
Spain
Definition
Synonyms
In chemistry, combustion is the complete or
Nucleic acid pool of random sequence; Oligonu- incomplete oxidation of a fuel by oxygen or
cleotide library other oxidants to give oxidized carbon species,
500 Comet
such as CO (for incomplete combustion) and CO2 meeting extreme physical conditions and as key
(for complete combustion), water and the con- objects for understanding the history of the Solar
comitant production of heat. For example, System.
The historical background of comets is
CH4 þ 2O2 ! CO2 þ 2H2 O reviewed in the books by Yeomans (1991) and
Schechner Genuth (1997). Important milestones
Respiration is an example of a biological com- for the science of comets were:
bustion reaction.
• The understanding that comets are not atmo-
spheric phenomena, which was demonstrated
See Also by Tycho Brahe (1546–1601) and his col-
leagues who measured the parallax of the
▶ Oxidation comet of 1577
▶ Respiration • The determination of cometary orbits with the
works of Isaac Newton (1644–1727) and
Edmond Halley (1656–1742)
• The knowledge of the comet phenomenon,
formation and development of both the coma
Comet and the tails (end of the eighteenth and nine-
teenth centuries)
Jacques Crovisier • The determination of the composition of
LESIA, Observatoire de Paris, Meudon, France comets with the emergence of spectroscopy
(end of the nineteenth century) and the open-
ing of new spectral domains (end of the twen-
Keywords tieth century)
• The active exploration of comets with flyby
Small body; Coma and encounter space missions nowadays
Cometary science is the topic of several books

Definition and reviews, such as those of Festou et al. (1993a, b,
2004) and Krishna Swamy (2010).
A comet is a small body, formed in the outer
region of the Solar System, generally on a highly
eccentric orbit, and containing a large fraction of Overview
volatiles. When coming close to the Sun, ▶ sub-
limation of the ices causes the development of Names of Comets
spectacular cometary phenomena: the coma and The nomenclature system for comets has changed
the dust and ion tails. several times. The Central Bureau for Astronom-
ical Telegrams (CBAT) of the International Astro-
nomical Union (IAU) is in charge of attributing
History names to comets. A registration code is given
according to the order of discovery. It consists of
Comets are among the most remarkable sky phe- “C/” followed by the year of discovery, then by a
nomena for both the layman and the scientist letter corresponding to the half-month of the dis-
(Fig. 1, Table 1). For a long time, their unex- covery, and then by an order numeral. In addition,
pected apparitions and their unknown nature the name of the discoverer (or of the first two
induced both fascination and fear. Nowadays, discoverers) is traditionally associated. Thus,
they are considered as natural laboratories C/1995 O1 (Hale-Bopp) is the first comet
Comet 501
Comet, Fig. 1 Comet

C/2006 P1 (McNaught)
over the Pacific. This comet
became spectacular as it
passed perihelion at
0.17 AU from the Sun on
12 January 2007 (#
Sebastian Deiries (ESO)) C
discovered in the second half of July 1995, by The inspection of the Catalog of Cometary
Alan Hale and Thomas Bopp. Short-period comets Orbits shows that there are two main families of
with a period <200 years are registered “P/” comets (e.g., Morbidelli 2008):
instead of “C/.” Those that have been observed at
several returns are given subsequently a number: • Short-period comets with low inclination over
1P/Halley, 2P/Encke, etc. See http://www.cfa.har- the ecliptic. Most of them have orbital periods
vard.edu/iau/lists/CometResolution.html for more close to 6 or 12 years. Their orbital evolution
details, exceptions, and oddities. is governed by strong gravitational interaction
One should be careful to comply with these with Jupiter. They are named “ecliptic
IAU rules to avoid ambiguities. In particular, comets” or “Jupiter-family comets.”
using only the discoverer(s) name(s), as is often • Comets with random inclination over the
the case in popular articles and even in some ecliptic. They may have a long orbital period
professional papers, should be discouraged (“new” comets with a nearly parabolic orbit)
(some discoverers were very prolific). or a short period (such as Halley’s comet).
They are named “nearly isotropic comets.”
Cometary Orbits, Cometary Families, and To explain the continuing supply of new
Cometary Reservoirs comets, the existence of a distant, spherical
There are up to now more than 3,708 cometary reservoir of comets was postulated (▶ Oort
apparitions for which secure the orbits and could Cloud). Thus, these comets are alternatively
be determined as listed in the Catalog of Come- named “Oort-cloud comets.”
tary Orbits (Marsden and Williams 2008). As of
June 2013, 281 comets have been observed at Nearly isotropic comets are not believed to
multiple returns and are now “numbered” short- have formed in the Oort Cloud but in the tradi-
period comets. tional Solar System. They were subsequently
Some comets have slightly hyperbolic orbits. ejected to the Oort Cloud following gravitational
As far as this could be investigated, they just interaction with giant planets. In turn, ecliptic
recently underwent gravitational perturbations comets could have formed in the trans-Neptunian
with a planet (Jupiter in most of the cases) region, among the ▶ Kuiper Belt. They subse-
which changed their orbit from elliptic to quently evolved to shorter-period orbits, preserv-
hyperbolic. The apparition of a genuine interstel- ing their low inclination.
lar comet, expelled from an extrasolar system, is Also related to comets are Centaurs, which are
not ruled out, but we are still awaiting for such an intermediate objects between Kuiper-Belt objects
event. and main-belt asteroids. Some of them, like
502 Comet
Comet, Table 1 A selection of well-known comets

(a) (b) (c) (d) (e) (f) (g) (h)
C/1577 V1 1577 I 27 Oct. 1577 0.178 1.0 First parallax
measurement
C/1680 V1 “Kirch’s 1680 18 Dec. 1680 0.0062 1.000 Sungrazer
comet”
C/1729 P1 1729 16 Jun. 1829 4.05 1.0 Intrinsically very
“Sarabat’s comet” bright
C/1743 X1 1744 1 Mar. 1744 0.222 1.0
“Chézeaux’s comet”
D/1770 L1 Lexell 1770 I 14 Aug. 1770 0.67 0.786 5.6 Approached
Earth at 0.015 AU
C/1811 F1 Great 1811 I 12 Sep. 1811 1.04 0.995
comet
C/1819 N1 Great 1819 II 28 Jun. 1819 0.342 1.0 First polarization
comet observation
C/1843 D1 Great 1843 I 27 Feb. 1843 0.0055 0.999 513 Sungrazer
March comet
C/1858 L1 Donati 1858 VI 30 Sep. 1858 0.578 0.996 2,000
C/1861 J1 Great 1861 II 12 Jun. 1861 0.82 0.985 409
comet
C/1864 N1 Tempel 1864 II 16 Aug. 1864 0.91 0.996 First spectral
C/1868 L1 1868 II 26 Jun. 1868 0.58 1.0 observations
Winnecke
C/1874 H1 Coggia 1874 III 1874c 9 Jul. 1874 0.68 0.998
C/1881 K1 Great 1881 III 1881b 16 Jun. 1881 0.73 0.996
comet
C/1882 R1 Great 1882 II 1882b 17 Sep. 1882 0.00775 0.9999 759 Sungrazer
September comet
C/1887 B1 Great 1887 I 1887a 11 Jan. 1887 0.0048 1.0 Sungrazer
southern comet
C/1901 G1 Great 1901 I 1901a 24 Apr. 1901 0.245 1.0
comet
C/1907 L2 Daniel 1907 IV 1907d 4 Oct. 1907 0.512 0.999
C/1908 R1 1908 III 1908c 26 Dec. 1908 0.945 1.0007
Morehouse
C/1910 A1 Great 1910 I 1910a 17 Jan. 1910 0.129 0.9999
January comet
C/1911 O1 Brooks 1911 V 1911c 28 Oct. 1911 0.49 0.997
C/1927 X1 1927 IX 1927k 18 Dec. 1927 0.176 0.9998
Skjellerup-
Maristany
C/1940 R2 1941 I 1940c 16 Jan. 1941 0.368 1.0005
Cunningham
C/1947 X1 Southern 1947 XII 1947n 2 Dec. 1947 0.110 0.9995
comet
C/1948 V1 Eclipse 1948 XI 1948l 27 Oct. 1948 0.135 0.99994
comet
C/1956 R1 Arend- 1957 III 1956h 8 Apr. 1957 0.316 1.0002
Roland
(continued)
Comet 503
Comet, Table 1 (continued)

(a) (b) (c) (d) (e) (f) (g) (h)
C/1957 P1 Mrkos 1957 V 1957d 1 Jul. 1957 0.355 0.9994
C/1969 Y1 Burnham 1960 II 1959k 29 Mar. 1960 0.355 0.9994
C/1961 R1 1962 VIII 1961e 10 Dec. 1962 2.13 0.990
Humason
C/1965 S1 Ikeya- 1965 VIII 1965f 21 Oct. 1965 0.00779 0.99992 Sungrazer C
Seki
C/1969 Y1 Bennett 1970 II 1969i 20 Mar. 1970 0.538 0.996
C/1973 E1 1973 XII 1973f 28 Dec. 1973 0.142 1.000 International obs.
Kohoutek campaign
C/1975V1 West 1976 VI 1975n 25 Feb. 1976 0.197 1.000
C/1980 E1 Bowell 1982 I 1980b 12 Mar. 1982 3.364 1.057 Active far from
the Sun
C/1983 J1 Sugano- 1983V 1983e 1 Apr. 1983 0.471 1.000 Approached
Saigusa-Fujikawa Earth at 0.063 AU
C/1983 H1 IRAS- 1983 VII 1983d 21 May 1983 0.991 0.990 Approached
Araki-Alcock Earth at 0.031 AU
C/1983 O1 Cernis 1983 XII 1983l 21 Jul. 1983 3.33 1.002 Active at more
C/1986 P1 Wilson 1987 VII 1986l 20 Apr. 1987 1.200 1.0003 than 20 AU
C/1989 X1 Austin 1990 V 1989c1 10 Apr. 1990 0.350 1.0002
C/1990 K1 Levy 1990 XX 1990c 24 Oct. 1990 0.939 1.0004
D/1993 F2 91994X 1993e Crashed on
Shoemaker-Levy Jupiter
C/1996 B2 1 May 1996 0.230 0.999 8,9000 Approached
Hyakutake Earth at 0.10 AU
C/1995 O1 Hale- 1 Apr. 1997 0.914 0.995 2,400 Extensive obs.
Bopp campaign
C/1999 S4 LINEAR 26 Jul. 2000 0.765 0.9994 Nucleus broke at
perihelion
C/2001 A2 LINEAR 24 May 2001 0.779 0.9993 Successive
C/2002 T7 LINEAR 23 Apr. 2004 0.615 1.0006 nucleus breakings
C/2001 Q4 NEAT 15 May 2004 0.962 1.0006
C/2004 Q2 24 Jan. 2005 1.205 0.9995
Machholz
C/2006 P1 12 Jan. 2007 0.171 1.0000 Reached
McNaught mv = 5 at
perihelion
Some numbered short-period comets
(a) (d) (e) (f) (g) (h)
1P/Halley 9 Feb. 1986 0.587 0.967 76.0 Target of several space missions
2P/Encke 9 Sep. 2000 0.340 0.847 3.30 Comet of shortest period
3D/Biela 23 Sep. 1852 0.861 0.756 6.62 Split comet now lost
9P/Tempel 1 5 July 2005 1.505 0.518 5.51 Target of Deep Impact mission
17P/Holmes 4 May 2007 2.053 0.432 6.88 Huge outburst in October 2007
19P/Borrelly 14 Sep. 2001 1.358 0.624 6.86 Target of Deep Space 1 mission
21P/Giacobini-Zinner 5 Sep. 1985 1.028 0.708 6.59 Target of ICE space mission
26P/Grigg-Skjellerup 22 July 1992 0.995 0.664 5.10 Target of Giotto extended mission
(continued)
504 Comet
Comet, Table 1 (continued)

Some numbered short-period comets
(a) (d) (e) (f) (g) (h)
29P/Schwassmann-Wachmann 1 10 July 2004 5.724 0.044 14.7 Nearly circular orbit
55P/Tempel-Tuttle 28 Feb. 1998 0.977 0.906 33.2 Assoc./Leonid meteor stream
67P/Churyumov-Gerasimenko 12 Aug. 2015 1.243 0.641 6.45 Target of Rosetta mission
73P/Schwassmann-Wachmann 3 7 June 2006 0.939 0.693 5.36 Split comet
81P/Wild 2 25 Sep. 2003 1.590 0.539 6.40 Target of Stardust mission
95P/Chiron 14 Feb. 1996 8.454 0.383 50.7 Centaur, also listed as asteroid
103P/Hartley 2 28 Oct. 2010 1.059 0.695 6.47 Target of EPOXI mission
109P/Swift-Tuttle 12 Dec. 1992 0.958 0.964 133.3 Assoc./Perseid meteor stream
133P/Elst-Pizarro 29 June 2007 2.642 0.164 5.61 Main-belt comet
(a) Official IAU designation
(b) Old-style designation before 1995
(c) Provisional designation before 1995
(d) Date of perihelion
(e) Perihelion distance [AU]
(f) Eccentricity
(g) Period (years)
(h) Remarks
Chiron (which is registered both as asteroid and fragile bodies (see Lamy et al. 2004;
(2060) Chiron and comet 95P/Chiron), show Weissman et al. 2004 and the ▶ Comet
cometary activity. (Nucleus) entry).
“Main-belt comets” are main-belt asteroids Water ice sublimates in the vacuum at temper-
which occasionally show a low level of cometary atures greater than about 150 K. For cometary
activity. They are also known as “activated aster- nuclei, this occurs at distances smaller than about
oids.” About half a dozen of such objects (133P/ 4 AU from the Sun; this results in the develop-
Elst-Pizarro) have been recently identified. Their ment of a cometary atmosphere: the gaseous
activity might be triggered by impacts from other coma. The gravity of the comet nucleus is too
small bodies. small to retain this atmosphere, which expands
“Sungrazing comets” pass within a few solar with a velocity ranging from 0.5 to a few km/s,
radii from the Sun and are generally only detect- depending on the distance to the Sun and the gas
able at that moment. More than 1,900 objects production rate. Thus, the gas density drops rap-
have been registered as sungrazing comets from idly with increasing distance from the nucleus,
observations with space coronagraphs (SOHO or and the flow becomes collision-free. The inner
STEREO). Most of these bodies are meter-sized collisional region has a size of a few hundred to a
objects that do not survive after perihelion. few thousand kilometers, depending on the comet
outgassing. As in laboratory molecular flows, the
The Nature of Comets and Basic Cometary gas temperature drops rapidly: temperatures in
Processes the range 10–100 K are typically observed.
We now know that comet nuclei are solid icy The production rate of water at a distance of
conglomerates and that the sublimation of ices about 1 AU from the Sun is typically 1028 mole-
in comet nuclei is the motor of cometary activity, cules per second (300 kg/s) for small short-period
following the popular model of the “dirty snow- comets such as 9P/Tempel 1 and 67P/
ball” of Fred Whipple (1906–2004). Comet Churyumov-Gerasimenko, both the targets
nuclei are kilometer-sized low-density porous of space missions. It was 1030 molecules
Comet 505
s1 (30 t/s) for 1P/Halley and as large as 1031 mol- the infrared satellites IRAS and ISO. When the
ecules s1 (300 t/s) for the giant comet C/1995 Earth encounters such a cometary trail, a meteor
O1 (Hale-Bopp). shower may be observed (the link between
Comets can still be active at distances larger comets and meteor streams was first established
than 4 AU from the Sun, where water sublimation for the case of the Perseid meteors and comet
is inefficient. This situation requires the sublima- 109P/Swift-Tuttle).
tion of more volatile species, such as CO, CO2, or Cometary ions in the coma are accelerated C
CH4, that are responsible for the cometary activ- through magnetohydrodynamic interaction with
ity. Indeed, the production of carbon monoxide the solar wind to velocities of several hundred km
was observed in ▶ comet Hale-Bopp at distances s1. They form a thin straight tail, easily distin-
as far as 14 AU. guishable from the broad, curved dust tail. This
The dominant chemical process in cometary tail is mainly composed of CO+ and H2O+ ions. It
atmospheres is the progressive molecular photol- is remarkable that this interaction was proposed
ysis by the solar UV radiation. The lifetime of the by Ludwig Biermann (1907–1986) and modeled
water molecule at 1 AU from the Sun is about by Hannes Alfvén (1908–1995) before the solar
1 day, but it may be significantly shorter for wind was actually observed by space probes.
complex, organic molecules. Two-body reactions Atoms and molecules that undergo fluores-
are inefficient because molecules spend only a cence excited by the Sun are also accelerated
short time in the collision region where tempera- away from the Sun. The process is especially
ture is low and where the fraction of reactive ions efficient for the resonant D lines of sodium at
and radicals is still low. 589 nm. A neutral sodium tail results, which
The primary radiation mechanisms for mole- was peculiarly conspicuous in ▶ comet Hale-
cules, radicals, atoms, and ions are the ▶ fluores- Bopp (Cremonese et al. 1997). The same process
cence of their electronic and vibrational bands is responsible for a significant distortion of the
excited by solar radiation and thermal emission large hydrogen coma.
of rotational lines. Prompt (non-fluorescence)
emission from radicals and atoms, following Space Missions to Comets
their creation in an excited state as a result of A table listing past and current space missions to
photolysis, may also occur. For rotational lines comets is given in the ▶ Comet Nucleus entry.
of molecules in the radio and far-infrared The main steps of cometary exploration were
domains, thermal excitation by collisions with (Keller et al. 2004):
molecules, ions, and electrons is another impor-
tant mechanism. • The ▶ VEGA and ▶ Giotto missions which
Cometary grains are dragged from the nucleus flew by 1P/Halley in March 1986 revealed
by gas. Their initial velocity is much smaller than the reality of a solid comet nucleus.
that of the gas and depends on the grain size. The • The ▶ Stardust mission which flew through
biggest “boulders” cannot escape the nucleus, the coma of 81P/Wild 2 on 2 January, 2004,
which contributes to the formation of a regolith. sampled cometary grains and returned them to
Dust grains are repelled by the Sun, forming the Earth for analysis on 15 January 2006.
sometimes spectacular dust tail, following the • The ▶ Deep Impact mission, an active exper-
kinematic model first proposed by Bessel iment, sent an impactor to the nucleus of
(1784–1846) and Bredichin (1831–1904). The 9P/Tempel 1 on 4 July 2005. The plume
physical process at work – the solar radiation which developed after the impact was
pressure – was later explained by Svante Arrhe- observed from the spacecraft and from the
nius (1859–1927). The biggest dust grains Earth. The same spacecraft, in a mission
migrate along the cometary orbit where they renamed EPOXI, explored 103P/Hartley 2 on
form cometary trails that were first imaged by 4 November 2010.
506 Comet
• The ▶ Rosetta mission (Schulz et al. 2009) mass spectroscopy. We now have a confident
will encounter ▶ 67P/Churyumov- knowledge of the main constituents of cometary
Gerasimenko in 2014–2015. An orbiter will ices (Bockelée-Morvan et al. 2004; Crovisier
stay in the comet vicinity for months, 2004; Crovisier et al. 2004; Fig. 2; Table 2).
witnessing the development of cometary The main components are water (about 80 % by
activity. A lander will make in situ analyses number), followed by carbon monoxide and diox-
of the nucleus material. ide. Then come methanol, ammonia, methane
and other hydrocarbons (C2H2, C2H6), hydrogen
An inescapable further step will be to return to sulfide, and hydrogen cyanide. About 15 other
Earth a sample directly taken from a comet minor constituents were identified in small
nucleus. Such missions are currently under amounts, but they can only be observed in the
study but are not yet firmly scheduled. most productive comets. For instance, the rela-
Space missions are not yet versatile enough to tively complex organic molecules formic acid
go to unexpected comets. Short-period comets, (HCOOH), methyl formate (HCOOCH3), acetal-
which have predicted returns, are presently the dehyde (CH3CHO), and ethylene glycol
only practicable targets. Flybys at low velocity (CH2OHCH2OH) have been identified by their
and rendezvous are only possible for ecliptic radio lines only in C/1995 O1 (Hale-Bopp).
(Jupiter-family) comets, due to energy limita- Indeed, the relationship between comets and
tions. Up to now, the only comet not belonging spectroscopy is exemplary and dates back from
to the Jupiter family that was explored was the beginning of astrophysics in the second half
1P/Halley; this was done with a very high flyby of nineteenth century. Many molecular species
velocity (about 70 km/s). were observed in comets before they could be
Although in situ cometary explorations are studied in the laboratory.
invaluable, they were (and will probably be for Mass spectroscopic measurements were
a long time) restricted to a very small number of performed in situ on 1P/Halley with Giotto
targets. Thus, a study of the comet diversity, (Altwegg et al. 1999, and references therein).
which needs a statistical approach, can only be However, their interpretation was hampered by
achieved with remote-sensing long-term observ- the limited mass resolution and the need for
ing programs. detailed chemical modeling to deduce neutral
abundances from the mass spectra. This problem
Composition of Comets: Ices and Volatiles of mass ambiguity will be resolved with
For a long time, the chemical study of comets was the equipment of the Rosetta orbiter. Also, the
restricted to visible spectroscopy. Radicals, Rosetta lander will directly investigate the
atoms, and ions, such as CN, CH, C2, C3, NH, nucleus material, with mass spectroscopy and
NH2, OH, CO+, etc. (Feldman et al. 2004), were gas chromatography.
observed. It was proposed in the mid-twentieth The ▶ Stardust mission returned samples from
century by Karl Wurm (1899–1975) and Pol comet 81P/Wild 2. The collecting technique
Swings (1906–1983) that these unstable species which was used (cometary grains were trapped
were “daughter molecules” coming from the by entering an aerogel substrate with a velocity of
photodestruction or photoionization by the solar 6 km/s) could not preserve volatiles and favored
UV radiation of volatile stable molecules, the refractory material. However, a careful analysis
“parent molecules,” released from the sublima- revealed the presence of methylamine
tion of nucleus ices. Proposed parents were H2O, (CH3NH2), ethylamine (CH3CH2NH2), and pos-
NH3, CH4, CO, CO2, HCN, etc. that could not be sibly glycine (NH2CH2COOH) in the returned
directly identified by the techniques available at aerogel that could be of cometary origin (Elsila
that time. et al. 2009). These species could result from
Confirmation came with the advent of radio, the degradation of carbonaceous cometary grains
infrared, and UV spectroscopy as well as in situ rather than from the sublimation of nucleus ices.
Comet 507
Comet, Fig. 2 Relative

production rates of H2O
cometary volatiles and their CO >10
comet-to-comet variations. CO2 7
These rates are believed to
trace the relative
CH4 8
abundances in cometary
ices. The red part of each C2H2 5
bar indicates the range of
C
C2H6 8
variation from comet to
comet. The number of CH3OH >10
comets in which the species
H2CO >10
was detected is indicated on
the right (Adapted from HOCH2CH2OH 1
Bockelée-Morvan HCOOH 3
et al. 2004)
HCOOCH3 1
CH3CHO 1
NH2CHO 1
NH3 3
HCN >10
HNCO 4
HNC 10
CH3CN 8
HC3N 2
H2S >10
OCS 2
SO2 1
CS2 >10
H2CS 1
NS 1
S2 5
10−3 10−2 10−1 1 101 102

Relative abundances (% relative to water)
Many features detected in cometary spectra at Indeed, from the observations of daughter spe-
all wavelengths are still unidentified, suggesting cies, A’Hearn et al. (1995) have identified a class
that new cometary species are still to be identi- of carbon-poor comets, for which the C2 radical is
fied. This requires further theoretical and labora- depleted. It appears that these carbon-poor
tory spectroscopic studies. comets are mostly present among Jupiter-family
Comets show a large diversity in their chem- comets. However, how this carbon depletion
ical composition (Fig. 2). For instance, the could be related to the abundance of bona fide
CO/H2O, CH3OH/H2O, or H2CO/H2O produc- parent molecules is still unclear. Clues from
tion rate ratios vary greatly from one comet to infrared and radio spectroscopy are yet inconclu-
another, whereas the HCN/H2O ratio is fairly sive, perhaps because the sample of investigated
constant. It is important to assess whether this comets at these wavelengths is still sparse
diversity is correlated with different sites of for- (Crovisier et al. 2009; Mumma and Charnley
mation or to different evolutions of these bodies. 2011; A’Hearn et al. 2012).
508 Comet
Comet, Table 2 The relative composition of volatiles observed in comet C/1995 O1 (Hale-Bopp), normalized to water
(From Bockelée-Morvan et al. 2004, with updates)
Water H2O 100
a
Carbon monoxide CO 12–23
Carbon dioxide CO2 6
Methane CH4 1.5
Acetylene C2H2 0.1–0.3
Ethane C2H6 0.6
Methanol CH3OH 2.4
a
Formaldehyde H2CO 1.1
Formic acid HCOOH 0.09
Methyl formate HCOOCH3 0.08
Acetaldehyde CH3CHO 0.02
Ethylene glycol CH2OHCH2OH 0.25
Formamide NH2CHO 0.015
Ammonia NH3 0.7
Hydrogen cyanide HCN 0.25
Isocyanic acid HNCO 0.10
a
Hydrogen isocyanide HNC 0.04
Methyl cyanide CH3CN 0.02
Cyanoacetylene HC3N 0.02
Hydrogen sulfide H2S 1.5
a
Carbonyl sulfide OCS 0.4
Sulfur dioxide SO2 0.2
b
Carbon disulfide CS2 0.2
Thioformaldehyde H2CS 0.05
c
NS radical NS 0.02
Hydrogen peroxide H2O2 <0.03
Propyne CH3CCH <0.045
Ketene CH2CO <0.032
Ethanol C2H5OH <0.10
Dimethyl ether CH3OCH3 <0.45
Glycolaldehyde CH2OHCHO <0.07
Acetic acid CH3COOH <0.06
Glycine I NH2CH2COOH <0.15
Cyanodiacetylene HC5N <0.003
Ethyl cyanide C2H5CN <0.01
Methenamine CH2NH <0.032
Cyanamide NH2CN <0.004
Methyl mercaptan CH3SH <0.05
Data derived from radio or infrared observations of comet Hale-Bopp made at heliocentric distances of about 1 AU
a
With possibly an additional distributed source in the coma
b
From the observation of the CS radical
c
Of unknown origin
Composition of Comets: Dust and (Semi-) observatories) and from the samples collected in
Refractories the coma of the Jupiter-family comet 81P/Wild
Knowledge of the composition of cometary dust 2 by the ▶ Stardust mission, providing ground
stems from infrared spectroscopy (with ground- truth for the remote-sensing investigations
based telescopes and the ISO and Spitzer space (Hanner and Zolensky 2010). Additional
Comet 509
information comes from the analysis of near-infrared spectroscopy (from features near
interplanetary dust particles (IDPs), collected in 3.4, 6, 8, and 12 mm) is subject to debate. More
the upper Earth’s atmosphere, which could be of compelling evidence of their presence comes
cometary origin. The analysis of the material from mass spectroscopy of the Stardust samples.
excavated in the Jupiter-family comet 9P/Tempel Icy grains released from the nucleus could be a
1 by the ▶ Deep Impact experiment revealed an significant source of gaseous material. At short
inner nucleus composition similar to that heliocentric distances, such grains have short C
observed in more active Oort-cloud comets, lifetimes, and their outgassing is difficult to dis-
such as 1P/Halley or C/1995 O1 (Hale-Bopp). tinguish from the nucleus outgassing with
Cometary dust appears to be heterogeneous, ground-based observations.
with silicates in both the amorphous (glassy) and
crystalline forms, Fe and Ni sulfides, and other Isotopic Ratios
minerals in minor amounts. The presence of car- Isotopic ratios and especially the D/H ratio are
bonates and phyllosilicates is subject to debate. important for investigating the circulation of mat-
Cometary silicates, which constitute the most ter in the Universe and in the Solar System (Robert
abundant part of the refractory grains, show a et al. 2000). The D/H ratio is a key parameter for
large diversity, comprising forsterite (Mg2SiO4), understanding the origin of water on Earth. The
enstatite (MgSiO3), olivines, and pyroxenes with first measurements of cometary D/H were
a wide range in Mg/Fe. Among the 81P/Wild restricted to a limited number of objects (1P/Hal-
2 samples were found highly refractory calcium ley, C/1995 O1 (Hale-Bopp), C/1996 B2
aluminum-rich inclusions (▶ CAIs) and frag- (Hyakutake)), all Oort-cloud comets (Jehin
ments of ▶ chondrules, as is usually found in et al. 2009). These D/H values clustered around
primitive meteorites. 3 104, which is about 12 times higher than the
A significant fraction of cometary dust is in the protosolar D/H value, but significantly below the
form of carbonaceous grains (also known as D/H ratio in some molecular species in dense
“CHON particles”), first found in the space explo- interstellar ▶ molecular clouds. In Earth’s oceans,
ration of comet Halley. They could be a potential the D/H is 1.5 104. However, recent observa-
source of molecules, alternative to the sublimation tions with the Herschel Space telescope indicate a
of volatiles from nucleus ices. They have been value close to the Earth’s ocean value in the
invoked to explain the distributed sources of Jupiter-family comet 103P/Hartley 2 (Hartogh
H2CO, CO, OCS, HNC, CN, etc. through et al. 2011) and an intermediate value for another
photodegradation or, more likely, thermal degra- Oort-cloud comet (Bockelée-Morvan et al. 2012).
dation (Cottin and Fray 2008). Polyoxymethylene This points to an isotopic diversity in the comet
(H2CO)n (POM), hexamethylenetetramine population (Fig. 3) and reopens the question on the
C6H12N4 (HMT), HCN polymers (HCN)n, and origin of terrestrial water.
carbon suboxide polymers (C3O2)n have been pro-
posed as cometary analogs, and their degradation Formation and Evolution of Cometary Matter
was studied in the laboratory. It is likely, however, The composition of cometary volatiles is strik-
that the real cometary carbonaceous grains have a ingly similar to that of some interstellar molecu-
more complex composition than these analogs, lar clouds (Table 3), especially of hot molecular
similar to the insoluble organic matter found in cores. This indicates that interstellar molecules in
the matrix of some ▶ carbonaceous chondrites. these clouds could come from the evaporation of
▶ Polycyclic aromatic hydrocarbon molecules comet-like ices. This could also suggest, as was
(PAHs), which are ubiquitous in the interstellar advocated by Mayo Greenberg (1922–2001), that
medium, are also expected to be present in cometary material comes from the direct agglom-
comets (see the review by Li 2009 and references eration of unprocessed interstellar grains. How-
therein). The identification of cometary PAHs ever, this is hardly tenable for cometary ices now,
from remote sensing, using near-ultraviolet or in view of our current understanding of the early
510 Comet
Hale− Bopp
Hyakutake
Enceladus
Ikeya−Zhang
Tuttle
10−3
2002T7
Halley
2009P1
Hartley 2
D/H ratio
Earth
10−4
Protosolar
ISM
10−5
Cl J S U N Oort cloud Jupiter family
Comet, Fig. 3 The D/H ratio observed in cometary water displayed for planets, the interstellar medium (ISM), and
compared to values in carbonaceous meteorites (CI), the the protosolar nebula refer to molecular H2 (From Hartogh
giant planets, the Earth’s oceans, and Enceladus. Data et al. 2011; Bockelée-Morvan et al. 2012)
Comet, Table 3 The relative compositions of interstellar Observations, confirmed by the results from
and cometary ices (From Despois and Cottin 2005) the ▶ Stardust mission, have shown that come-
Interstellar tary matter includes ices as well as crystalline and
Interstellar ices amorphous silicates, originating from the cold
ices high- low-mass Cometary and hot regions of the solar nebula. This points
Species mass YSOa YSOb volatilesc
to an important turbulent mixing of the nebula.
H2O 100 100 100
Thus, cometary silicates differ from interstellar
CO 9–16 6–25 1.7–23
silicates which are always in the amorphous state.
CO2 14–20 15–22 6
CH4 2 <1.6 0.6
Cometary refractory grains could have both an
C2H6 <0.4 – 0.6 interstellar and a nebular origin (Wooden 2008;
CH3OH 5–22 <4 0.9–6.2 A’Hearn 2011).
H2CO 1.7–7 – 0.13–1.3
HCOOH 0.4–3 – 0.09 Comets and the Origin of Life
NH3 13–15 <9 0.7 The possible role of comets as a vector for
X-CN 1–3 <0.4 0.08–0.25 ▶ panspermia was advocated in several papers,
OCS, 0.05–0.3 <0.08 0.4 especially by Hoyle and Wickramasinghe (e.g.,
XCS 1981). The objections raised to this hypothesis
a
High-mass Young stellar object such as W33A and are the hostile radiation environment and the easy
N753S
b destruction of organic material following impact
Low-mass Young stellar object such as Elias 16 and Elias
26 (a category to which the protosun belonged) on Earth.
c
From Table 1 and Fig. 2 Could liquid water exist in comets, favoring
the development of life in these bodies? As was
suggested by Irvine et al. (1980) and studied in
Solar System history. The similarity between detail by Podolak and Prialnik (2006), a signif-
interstellar and cometary matter would rather be icant pressure together with a heating source
due to a similarity of the physicochemical pro- is needed for this to be possible. The latter
cesses occurring in dense interstellar clouds and could be provided internally by 26Al radioactiv-
in the early solar nebula. ity or by amorphous crystalline water ice
Comet 511
transition. However, this would only be imagin- Barucci MA, Boehnhardt H, Cruikshank DP, Morbidelli
able inside huge cometary nuclei (>100 km) or A (2008) The solar system beyond Neptune. The Uni-
versity of Arizona Press, Tucson
large KBOs. Bockelée-Morvan et al (2012) Herschel measurements of
The hypothesized delivery of organic mole- the D/H and 16O/18O ratios in water in the Oort-cloud
cules to the Earth by the infall of small Solar comet C/2009 P1 (Garradd). Astron Astrophys 544:
System bodies was prompted by the discovery L15
Bockelée-Morvan D, Crovisier J, Mumma MJ, Weaver
of carbonaceous matter in meteorites in the nine- HA (2004) The composition of cometary volatiles.
C
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out by Chamberlin and Chamberlin (1908) and by II. The University of Arizona Press, Tucson,
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Chamberlin TC, Chamberlin RT (1908) Early terrestrial
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and Cottin 2005; Thomas et al. 2006). Space Sci Rev 138:179–197
Cremonese G, Boehnhardt H, Crovisier J, Rauer H,
Fitzsimmons A, Fulle M, Licandro J, Pollacco D,
Tozzi GP, West RM (1997) Neutral sodium from
See Also comet Hale-Bopp a third type of tail. Astrophys
J 490:L199–L202
Crovisier J (2004) The molecular complexity of comets.
▶ Carbonaceous Chondrite In: Ehrenfreund et al (eds) Astrobiology: future per-
▶ Centaurs (Asteroids) spectives. Kluwer, Dordrecht, pp 179–203
▶ Comet (Nucleus) Crovisier J, Bockelée-Morvan D, Colom P, Biver N,
Despois D, Lis DC (2004) The composition of ices in
▶ Comet Halley
comet C/1995 O1 (Hale-Bopp) from radio spectros-
▶ Daughter Molecule, Comet copy, further results and upper limits on undetected
▶ Deep Impact species. Astron Astrophys 418:1141–1157
▶ Fluorescence Crovisier J, Biver N, Bockelée-Morvan D, Boissier J,
Colom P, Lis DC (2009) The chemical diversity of
▶ Giotto Spacecraft
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▶ Kuiper Belt Despois D, Cottin H (2005) Comets: potential sources of
▶ Meteorites prebiotic molecules for the early earth. In: Gargaud M,
▶ Molecular Cloud Barbier B, Martin H, Reisse J (eds) Astrobiology.
Springer, Berlin, pp 289–352
▶ Oort Cloud
Elsila JE, Glavin DP, Dworkin JP (2009) Cometary gly-
▶ Panspermia cine detected in samples returned by Stardust. Meteorit
▶ Parent Molecule, Comet Planet Sci 44:1323–1330
▶ Rosetta Spacecraft Feldman PD, Cochran AL, Combi MR (2004) Spectro-
scopic investigations of fragment species in the coma.
▶ Stardust Mission
In: Festou MC, Keller HU, Weaver H (eds) Comets
▶ Vega 1 and 2 Spacecraft II. The University of Arizona Press, Tucson,
pp 425–447
Festou MC, Rickman H, West RM (1993a) Comets.
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Keller HU, Weaver H (eds) Comets II. The University
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CNES/DSP/SME, Vétérinaire/DVM, Astro/
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Central Bureau for Astronomical Telegrams,
Cambridge Definition
Morbidelli A (2008) Comets and their reservoirs: current
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The experiment “COMET” flew actually three
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Facility), and 1999 (COMET-99) outside the sta-
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compounds on the primitive earth. Nature material is composed of grains with a size distri-
190:389–390 bution dominated by particles in the micron
Podolak M, Prialnik D (2006) The conditions for liquid
range. Very long collection times are required to
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evolution of life, 2nd edn. Springer, New York, or more). For instance, the experiment, Orbital
pp 303–314 Debris Collection Experiment (ODCE) spon-
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92:201–224 space station and collected for 18 months starting
Schechner Genuth S (1997) Comets, popular culture and in March 1996 (Fig. 1).
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Comet (Nucleus) 513
Borg J (2002) Extraterrestrial samples from low earth

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C
Comet (Nucleus)
Jacques Crovisier
LESIA, Observatoire de Paris, Meudon, France
COMET (Experiment), Fig. 1 The COMET hardware Keywords

before being placed outside the MIR space station (Photo
RKA/CNES)
Small body
specifically the collection of grains from the Definition

swarm of Leonids originated from the Tempel-
Tuttle comet. A cometary nucleus is the small-size (typically a
The various collectors made of ultrapure metal- few kilometers in diameter) solid body in the
lic foil or aerogel, a transparent silicon dioxide head of a comet whose activity provides the
material of very low density (0.06 g/cm3), source of the comet’s appearance.
came back to Earth by the end of the mission. On
the metallic collectors, the incident particles sub-
limate upon impact, leaving a crater, on the rims of Overview
which remnants of the particles can be found. The
aerogel collectors are designed to avoid destroying With typical sizes from about 1 km to about 50 km
the particles while impacting the surface. Penetrat- (as for the giant comet C/1995 O1 (Hale-Bopp)),
ing inside the aerogel, they slow down and are cometary nuclei are too small to be imaged by
trapped with little alteration. The metallic collec- Earth-based telescopes. This can only be done
tors show a very high density of large craters, due with space probes that encounter or flyby these
to particles larger than 20 mm attributed to the objects at small distances. This has been possible
Leonid swarm. Chemical identification was possi- up to now only for a restricted number of objects.
ble in one case, the EDS spectrum showing iden- These observations confirmed that cometary
tification of Mg, Si, Ca, and Fe. nuclei are really solid bodies (Fig. 1, Table 1).
Optical scanning of the aerogel collectors The unresolved comet nucleus can be
locates grains as small as a few microns and observed when the comet is far from the Sun
reveals their penetration track. In one of the two and inactive or when the instrumental resolution
aerogel collectors of COMET-99, two allows us to separate the nucleus emission from
populations of grains, with a mean diameter that of the dust coma. It is then possible to eval-
around 5 mm, have been found, with penetration uate the nucleus size and to investigate its rota-
tracks in two main directions. A dozen of these tion, using the same methods as those used for
grains have been extracted. asteroids (Lamy et al. 2005). Radar studies are
The experience gained during these experiments also possible for the nucleus of comets coming
was used preparing for the ▶ Stardust mission. close to the Earth.
514 Comet (Nucleus)
Cometary nuclei are very dark, with ▶ albedos rotation (e.g., they are tumbling rather than rotat-
in the range 0.02–0.06, which makes them the ing around a fixed axis).
darkest objects of the Solar System. Rotation The density of comet nuclei is still waiting for
periods range from a few hours to a few days. confident measurement. This will be done when it
Some nuclei seem to be in an excited state of will be possible to observe and measure the tra-
jectory perturbation of a space probe in the small
gravity field of a cometary nucleus. Indirect eval-
uations were performed using the effect of
nongravitational forces, caused by nucleus
outgassing, on cometary orbits (Weissman
et al. 2005). They all point to small densities in
the range 0.5–1.2 g cm3, suggesting that the
nuclei could be porous bodies.
Comet nuclei are frequently observed to split,
showing that they are bodies with weak tensile
strength, such as rubble piles.
See the ▶ Comets entry for a discussion of
composition and other details on comet nuclei.
See Also
▶ Albedo
▶ Comet
▶ Deep Impact
▶ EPOXI Mission
▶ Rosetta Spacecraft
Comet (Nucleus), Fig. 1 The nucleus of 9P/Tempel 1 as
seen by the Deep Impact probe. The nucleus average ▶ Stardust Mission
diameter is 6 km (# NASA/JPL-Caltech/UMD) ▶ Vega 1 and 2 Spacecraft
Comet (Nucleus), Table 1 Space missions which were able (or are on their way) to image cometary nuclei
Space
Mission agency Launch date Target Encounter date (a) (b)
VEGA 1 IKI 15 1P/Halley 6 Mar. 1986 (flyby) 8,890 km 79 km/s
Dec. 1984
VEGA 2 IKI 21 1P/Halley 9 Mar. 1986 (flyby) 8,030 km 77 km/s
Dec. 1984
Giotto ESA 2 Jul. 1985 1P/Halley 14 Mar. 1986 (flyby) 596 km 68 km/s
Deep Space NASA 24 Oct. 1998 19P/Borrelly 22 Sep. 2001 (flyby) 2,170 km 17 km/s
1
Stardust NASA 7 Feb. 1999 81p/Wild 2 2 Jan. 2004 (flyby) 237 km 6 km/s
NeXT ” ” 9P/Tempel 1 14 Feb. 2011 (flyby) 178 km 11 km/s
Rosetta ESA 2 Mar. 2004 67P/CG 2014–2015 (orb./land.)
Deep Impact NASA 12 Jan. 2005 9P/Tempel 1 4 Jul. 2005 (flyby/ 500 km 10 km/s
impact)
EPOXI ” ” 103P/Hartley 4 Nov. 2010 (flyby) 700 km 12 km/s
2
For flybys are listed (a) the closest distance to the target and (b) the relative velocity. 67P/CG: Churyumov-Gerasimenko
Comet Giacobini-Zinner 515
A’Hearn MF et al (2011) EPOXI at comet Hartley 2. ▶ Comets, History of

Science 332:1396–1400
Boehnhardt H (2005) Split comets. In: Festou MC, Keller
HU, Weaver H (eds) Comets II. The University of
Arizona Press, Tucson, pp 301–316
Festou MC, Keller HU, Weaver H (2004) Comets II. The C
University of Arizona Press, Tucson Comet Encke
Keller HU, Britt D, Buratti BJ, Thomas N (2005) In
situ observations of cometary nuclei. In: Festou MC, Therese Encrenaz
Keller HU, Weaver H (eds) Comets II. The University
of Arizona Press, Tucson, pp 211–222 LESIA, Observatoire de Paris - Section de
Lamy PL, Toth I, Fernandez YR, Weaver HA (2005) The Meudon, Meudon, France
sizes, shapes, albedos, and colors of cometary nuclei. In:
Festou MC, Keller HU, Weaver H (eds) Comets II. The
University of Arizona Press, Tucson, pp 223–264
Samarasinha NH, Mueller BEA, Belton MJS, Jorda Definition
L (2005) Rotation of cometary nuclei. In: Festou
MC, Keller HU, Weaver H (eds) Comets II. The ▶ Comet 2P/Encke was named after Johannes
University of Arizona Press, Tucson, pp 281–299 Encke who, in 1821, calculated its orbit on the
Verveka J et al (2013) Return to comet Tempel 1: over-
view of Stardust-NExT results. Icarus 222:424–435 basis of previous measurements made by Pons in
Weissman PR, Asphaug E, Lowry SC (2005) Structure 1818. He predicted its return in 1821 with an
and density of cometary nuclei. In: Festou MC, Keller accuracy of 1 day. In 1823, he identified that
HU, Weaver H (eds) Comets II. The University of comet Encke had already appeared in 1786,
Arizona Press, Tucson, pp 337–357
1795, and 1805. Comet Encke was thus the sec-
ond known periodic comet. Its period (3.3 years)
is the shortest reported until today. Its perihelion
is 0.33 AU. After over 60 passages since its first
discovery, comet Encke appears as a weak object
Comet Borrelly
that has lost most of its gas and dust.
Therese Encrenaz
See Also
▶ Comet
Definition
Comet 19/P Borrelly is a short-period comet,

discovered in 1904. Its orbital period is Comet Giacobini-Zinner
6.8 years. Its semimajor axis is 3.49 AU, and the
aphelion and perihelion distances to the Sun are Therese Encrenaz
5.83 and 1.35 AU, respectively. Comet Borrelly LESIA, Observatoire de Paris - Section de
was visited on September 22, 2001, by the space- Meudon, Meudon, France
craft Deep Space 1 that approached the comet at a
distance of about 2,200 km. The spacecraft
returned high-quality images of the comet Definition
nucleus, revealing the presence of several jets.
The comet is 8 4 4 km in size, and its den- ▶ Comet 21P/Giacobini-Zinner, discovered
sity is estimated as 0.3 g/cm3. Its albedo is 0.03, a independently by Michel Giacobini in 1900 and
typical value for comets. Ernst Zinner in 1913, is a Jupiter-family comet
516 Comet Hale–Bopp
with a period of 6.6 years; its perihelion is at Overview

1.0 AU and its aphelion is at 6.0 AU. In Septem-
ber 1985, the comet was visited by the spacecraft A large international observing campaign was
ISEE 3, renamed International Cometary devoted to the comet in 1996 and 1997, using
Explorer (ICE), which flew by the comet at a ground-based and space observatories at all
distance of 7,800 km. The spacecraft detected a wavelengths, from the X-ray to the radio range.
shock front when the ionized cometary species Visible cameras (in particular aboard the Hubble
came in contact with the solar wind. Six months Space Telescope) were used to determine the
before the encounter of comet Halley by five coma structure and the nucleus size and rotation.
spacecraft, 21P/Giacobini-Zinner was the first ▶ Parent molecules were mostly studied from
comet to be investigated by a space mission. infrared, millimeter, and submillimeter spectros-
copy, including space observations from the
Earth-orbiting Infrared Space Observatory (ISO).
See Also With a diameter of 40–80 km, the nucleus of
comet Hale–Bopp is the largest cometary nucleus
▶ Comet ever measured. Its rotation period, determined
from the evolution of the jets, is 11.4 h, a typical
value for cometary nuclei. In addition to the dust
and plasma tails usually visible on comets, a thin
Comet Hale–Bopp sodium-atom tail, observed in the visible range,
was detected for the first time (Fig. 1).
Therese Encrenaz Many parent molecules were detected through
LESIA, Observatoire de Paris - Section de infrared and millimeter spectroscopy. In addition
Meudon, Meudon, France to previously detected species (H2O, CO, CO2,
CH3OH, H2CO, HCN, H2S, NH3, HNCO,
CH3CN, HNC, and OCS), new molecules were
Keywords found in the radio range: HCOOH, CH3CHO,
HCOOCH3, NH2CHO, HC3N, H2CS, SO, SO2,
Comet and NS. The detection of HDO in the submillime-
ter range, in complement with the estimate of H2O
inferred from the radio monitoring, led to the
Definition determination of D/H. As for comets Halley and
Hyakutake, the D/H was found to be 3 104,
Comet C/1995 O1 (Hale–Bopp) was discovered i.e., twice its value in the terrestrial oceans; this
simultaneously by Alan Hale and Thomas result is important as it leads to the conclusion that
Bopp on July 23, 1995, at 7.1 AU from the Sun. only a minor fraction of water on Earth probably
It was soon realized that the comet, 100 times has a cometary origin. Ground-based observations
brighter than Halley at the same heliocentric dis- in the near-infrared range led to the detection of
tance, was abnormally big. Its 2,400-year period H2O, CO, CH4, C2H2, C2H6, OCS, NH3, as well as
brought it to perihelion on April 1, 1997, at a several radicals.
distance of 0.91 AU from the Sun. It was a Another important result is the determination
naked-eye object for over 2 months. As all long- of the nature of the cometary dust, inferred from
period ▶ comets, comet Hale–Bopp is an ▶ Oort ▶ Infrared Space Observatory measurements. In
cloud comet, one of the greatest of the twentieth addition to already known silicate signatures, the
century. Thanks to its early discovery and its spectrum of Hale–Bopp, recorded over the whole
exceptional size, comet Hale–Bopp is among the infrared range, exhibited specific signatures which
few objects which have allowed a major advance were attributed to forsterite, a magnesium-rich
in our understanding of cometary physics. olivine (Mg2SiO4). This spectrum is remarkably
Comet Halley 517
Comet Hale–Bopp, Fig. 1 Comet Hale–Bopp, observed from Earth during its 1997 passage (# M. Jourdain de
Muizon 1997)
similar to those of dust disks surrounding young or 76-year period, it is the only bright comet
evolved stars, showing a close similarity between whose trajectory is predictable enough for space
interstellar and cometary dust. exploration to be planned in advance. It is also the
comet which allowed the astronomer Edmund
Halley to demonstrate the nature and the periodic
See Also appearance of these objects: in 1705, on the basis
of previous observations, he predicted the
▶ Comet comet’s return in 1758. The comet’s apparition
▶ Oort Cloud was actually observed in December 1758,
▶ Parent Molecule, Comet 16 years after Halley’s death. The apparition of
comet Halley was decisive in confirming New-
ton’s laws of universal gravitation.
Comet Halley
Overview
Therese Encrenaz
LESIA, Observatoire de Paris - Section de Early apparitions of comet Halley go back to
Meudon, Meudon, France antiquity. Its 1066 CE apparition was recorded
in the Bayeux tapestry showing King Harold’s
fear shortly before the Hasting battle and the
Keywords victory of William the Conqueror. In 1301, the
famous Italian painter Giotto di Bondone
Comet represented the comet as the Bethlehem star in
his fresco “Adoration of the Magi” in Padua.
The 1835 comet Halley apparition led to the
Definition first observations of physical phenomena such as
gas and dust ejection in the form of jets and fans
Known since antiquity, comet Halley is probably around the nucleus, represented, in particular, in
the most famous comet in the world. With its Bessel’s drawings. His “fountain model” gave the
518 Comet Halley
first interpretation of the motion of cometary Comet Halley was recovered in October 1982,
material ejected sunward and being repelled at a heliocentric distance of 11 AU, with an
away from the Sun. This was the first confirma- angular distance of only 9 arcsec from its
tion of Laplace’s predictions, in 1803, about a predicted position. The observation was made at
frozen nucleus. The next apparition, 1910, was Mount Palomar Observatory (California, USA)
especially favorable in terms of geometrical con- and its recovery illustrated the growing success
figuration: photographs and spectra of the comet of ▶ CCD cameras in astronomy.
were recorded, showing emissions from several
radicals and ions. At that time, secondary prod- The Nucleus
ucts (coming from the photolysis and ionization The first images of a cometary nucleus were those
of parent molecules), observed in the visible of comet Halley, taken by the Giotto spacecraft in
range, were much better known than the parent 1986. They were surprising at least in two
molecules themselves which are better probed at aspects: the nucleus had an elongated shape,
infrared and millimeter wavelengths. In 1950, on with dimensions of 15 8 7 km in size, and
the basis of the H and OH production rates, the its mean albedo was very low (0.04). The nucleus
astronomer Fred Whipple proposed his “dirty surface was mostly covered with dark material,
snowball model,” which predicted that the probably due to a carbonaceous deposit, with a
▶ comets are mainly composed of water ice surface temperature as high as 300 K; water
together with minor amounts of dust. vapor, together with dust, was outgassed through
The next apparition, in 1986, was not favorable a few discrete active areas, at temperatures of
in terms of geometry: with respect to the Earth, the about 200–220 K (Fig. 1).
comet was behind the Sun at the time of perihelion, The presence of carbonaceous material at the
on February 9, 1986. Ground-based monitoring surface of the nucleus was also inferred by
was thus much more difficult than at the time of another discovery. The mass spectrometers of
the 1910 apparition, and the images of the comet the Giotto and Vega spacecraft detected
were much less spectacular; the best observations
were performed when the geocentric distance of
the comet was minimum in November 1985 and in
April 1986. In spite of the poor observational con-
ditions, the development of new technologies, both
for ground-based and space explorations, allowed
astronomers to get excellent scientific return from
this event. The apparition was prepared long in
advance by an impressive worldwide observing
campaign, including five spacecraft- and ground-
based observations at all wavelengths, from the
UV to the radio range, coordinated by the “Inter-
national Halley Watch.” The five space missions
included the European Giotto mission, launched
by the ESA, the two Vega missions led by the
Soviet Union, and two Japanese spacecraft, Suisei
and Sakigake. For the first time, images of the
nucleus were obtained; in situ measurements of
the cometary gas and dust were recorded; from Comet Halley, Fig. 1 The nucleus of comet Halley, as
space and ground-based experiments, parent mol- observed by the camera of the European Giotto mission on
March 13, 1986, as the spacecraft flew over the comet at a
ecules were actually detected, thanks to the devel-
distance of 500 km. Active areas showing jets of water and
opment of infrared spectrometers and millimeter dust appear on the left side of the object. The dimensions
heterodyne spectroscopy. of the nucleus are 15 8 7 km (# ESA)
Comet Halley 519
unexpectedly large abundances of light elements production of the comet as a function of its helio-
(H, C, N, O), especially in complex hydrocarbon centric distance.
grains. This result was independently derived An important parameter measured by mass
from the analysis of the near-infrared spectrum spectroscopy was the D/H ratio inferred from
of Halley recorded by Vega, which revealed a HDO/H2O in the coma. This ratio is diagnostic of
broad emission attributed to both saturated and the early conditions of the comet’s formation: at
unsaturated hydrocarbons. Such spectral signa- low temperatures, the D/H ratio is enriched in ices C
tures have been observed under other circum- as a result of ion-molecule and molecule-molecule
stances in interstellar spectra and also in reactions. D/H in comet Halley was found to be
laboratory spectra of ice mixtures irradiated by 3 104, i.e., twice its value in the terrestrial
solar UV or high-energy particles. oceans. This value was later confirmed by two
Long-period ground-based photometric moni- measurements on other comets (Crovisier 2001;
toring has been used to determine the rotation Newburn et al. 1999), which led to the same results.
period of Halley’s nucleus. A two-component If these comets are representative of most comets,
model has been favored with a slow precession the relatively high D/H ratio indicates that water on
(with a period of at least 7 days) and a 2-day Earth cannot have come entirely from comets.
rotation period.
Origin and Fate of Comet Halley
The Coma As shown by its retrograde orbit, comet Halley is
As a first result, Whipple’s “dirty snowball most likely a captured object from the Oort cloud.
model” was confirmed. Water, identified by its The ▶ Oort cloud is the reservoir to which many
near-infrared vibration bands, was unambigu- planetesimals were ejected by gravity perturba-
ously detected, both from the Kuiper Airborne tions due to the ▶ giant planets. This explains
Observatory and from the Vega IKS spectrome- why, after so many apparitions, comet Halley is
ter. The relative water content, in number of still an active body: its long period allowed it to
atoms, was found to be close to 80 %. In addition keep a significant fraction of its icy reservoir. At
to water and hydrocarbons, other molecules were each perihelion passage, the surface of comet
detected by IKS aboard Vega: CO2, H2CO, and Halley is eroded and looses about 1 m depth of
CO were also detected in the UV range. Hydro- ice and dust. In the future, the icy content of
gen cyanide HCN was detected for the first time comet Halley will slowly decrease.
in a comet from ground-based millimeter hetero-
dyne spectroscopy. Some molecules are
outgassed from the nucleus but some also from See Also
the grains in the coma. This is the case for CO and
H2CO, which have been detected in the cometary ▶ Comet
grains. Another interesting result was the ▶ Comet (Nucleus)
reported detection, by the UV spectrometer of ▶ Daughter Molecule, Comet
Vega, of some polycyclic aromatic hydrocarbons ▶ Giotto Spacecraft
(PAHs), probably naphthalene C10H8 and phen- ▶ Parent Molecule, Comet
anthrene C14H10, in the close vicinity of the ▶ Vega 1 and 2 Spacecraft
nucleus. Similar species have been also identified
in interstellar spectra. It is in fact striking to note
that all parent molecules found in comet Halley References and Further Reading
have been also detected in the interstellar
medium. These results strongly suggest a close Crovisier J (2001) Comet Hale-Bopp. In: Encyclopedia of
astronomy and astrophysics. Institute of Physics Pub-
link between interstellar and cometary matter.
lishing, Bristol, pp 414–416
Radio observations of the OH radical at wave- Newburn RL, Rahe J, Neugebauer M (1999) Comets in the
length of 18 cm were used to monitor the water post-Halley era. Kluwer, Dordrecht
520 Comet Hartley 2
March 1996 to a distance of only 0.10 AU. It

Comet Hartley 2 passed perihelion in May 1996 at a distance of
0.23 AU. It is a long-period comet (about
Therese Encrenaz 9,000 years) which presumably comes from the
LESIA, Observatoire de Paris - Section de ▶ Oort cloud. A campaign of astronomical obser-
Meudon, Meudon, France vations was set up, taking advantage of its prox-
imity to Earth. The comet nucleus is 2–3 km in
diameter and its rotation period is 6.3 h. Many
Definition parent molecules were detected from infrared and
millimeter ground-based spectroscopy (H2O,
Comet 103 P/Hartley 2, discovered in 1986, is a CO, CO2, CH3OH, H2CO, HCN, and H2S).
Jupiter-family comet with an orbital period of
6.46 years and a semimajor axis of 3.46 AU. Its
perihelion distance to the Sun is 1.05 AU, close to See Also
Earth’s orbit. In 2010, the comet approached the
Earth to a distance of 0.12 AU. Radar observa- ▶ Comet
tions revealed an elongated shape and measured a ▶ Oort Cloud
rotation period of 18 h. On November 2010, the
comet was visited by the ▶ Deep Impact space-
craft, renamed Epoxi, after its flyby of comet
Tempel 1. Epoxi approached the comet at a dis- Comet Mc Naught
tance of 700 km, measured its maximum size
(2.25 km), and revealed an unexpected activity. Therese Encrenaz
In 2011, the Herschel space observatory mea- LESIA, Observatoire de Paris - Section de
sured heavy water HDO in the coma and inferred Meudon, Meudon, France
a D/H ratio close to the terrestrial value. This
result suggests that the ocean water might come,
at least partly, from Jupiter-family comets. Definition
Over 50 comets have been discovered by Robert

See Also H. McNaught since 1987. Comet C/2006 P1
McNaught is a special comet which crossed peri-
▶ Comet helion at 0.17 AU from the Sun in December
2006 and became, in January 2007, the brightest
and largest comet observed from Earth for
40 years. Infrared observations from Earth led
to the identification of a large number of parent
Comet Hyakutake molecules. The unusually large size of the ion-
ized comet’s tail was explored as the Ulysses
Therese Encrenaz spacecraft passed through it in 2007. The comet
LESIA, Observatoire de Paris - Section de is a dynamically active ▶ Oort cloud comet, and
Meudon, Meudon, France its material is expected to be pristine, reflecting
early conditions of the solar system’s history.
Definition
See Also
Comet C/1996 B2 Hyakutake was discovered in
January 1996 and it approached the Earth in ▶ Comet
Comet Shoemaker-Levy 9 521
The fragments of SL9 entered the Jovian atmo-

Comet Shoemaker-Levy 9 sphere at a latitude of 44 south (44S). Between
July 16 and July 22, 1994, the impacts spread along
Therese Encrenaz the 44S parallel as the planet was rotating; the
LESIA, Observatoire de Paris - Section de collisional events took place within a few minutes
Meudon, Meudon, France of the predicted impact times. As seen from Earth,
the impacts took place just behind the limb and C
came to direct view only about 10 min later. The
Keywords Galileo spacecraft, still at a distance of 1.6 AU
from Jupiter, was the only observatory which
Comets could see the whole event in direct view.
The successive events were recorded with
three types of data: light curves, images, and
Definition spectra. Light curves, measuring the brightness
versus time at different wavelengths, allowed
Comet Shoemaker-Levy 9 (SL9) was discovered astronomers to determine the temperature evolu-
in March 1993 by Eugene and Carolyn Shoe- tion and gave an estimate of the energy budget.
maker and by David Levy. At that time, it Images, taken from Galileo, the HST, and
appeared as a trail of about 20 fragments. The ground-based telescopes, recorded the meteor
study of their trajectories showed that it was a (cometary fragment) entry (Galileo), the ejecta
Jupiter-family comet which was disrupted by trajectories (limb images from the HST), and the
tidal forces at its previous closest (perijove) pas- crater evolution (HST and ground-based tele-
sage, in July 1992. It was soon predicted that the scopes). Spectra, recorded in the UV (IUE,
fragments would collide with ▶ Jupiter at its next HST), the visible, infrared, and radio range,
closest passage, in July 1994 (Fig. 1). have detected new molecular species and moni-
tored their evolution.
Overview Temporal Sequence of the Impacts

The collisions of each individual fragment were
A collision of a comet with Jupiter is very rare: remarkably similar, although of different intensi-
according to dynamical models, it could be ties, depending on the impactor size. In all cases,
expected only once every few centuries. Actu- the light curves showed a sequence of three
ally, a similar event was reported by ▶ Cassini phases:
at the end of the seventeenth century. As a con-
sequence, at the time of the SL9 collision, a huge • The entry phase was observed as a flash by the
international campaign took place in order to camera and the photometer of the Galileo
monitor the event using all possible ground- spacecraft.
based and space means, covering the whole spec- • The explosion phase occurred 1 min later; it
tral range, from the X-ray to the radio range. corresponded to the explosion of the impactor
Among the spacecraft were the Galileo spacecraft within the atmosphere and to the ascent of a
(en route to Jupiter for an approach in 1995), the fireball; HST images taken at Jupiter’s limb
▶ Hubble Space Observatory (HST), the Interna- showed ejecta up to an altitude of 3,000 km for
tional Ultraviolet Explorer (IUE), and the X-ray all fragments (the smaller exploding at a
satellite ROSAT. Among the objectives were the higher altitude than the larger ones). The fire-
determination of the impact altitude, the temper- ball increased in size adiabatically from
ature rise and decay, the monitoring of the impact 15 km, 10 s after the impact, to 100 km after
craters, and the search for new molecules formed 40 s. Explosions took place at a pressure level
by shock chemistry. ranging between 0.1 and 1 bar. For the largest
522 Comet Shoemaker-Levy 9
Comet Shoemaker-Levy 9, Fig. 1 Image of the comet Shoemaker-Levy 9 before its impact with Jupiter (# NASA)
impacts, the temperature was over 10,000 K at • From mid-infrared spectra (10–13 mm), NH3,
the early beginning and decreased to 2,000 K HCN, and C2H4 were identified.
after about 15 s. • Ground-based millimeter heterodyne spec-
• The “splash phase” was studied from the light troscopy allowed the detection and long-term
curves, the images, and the evolution of the monitoring of CO, CS, OCS, and HCN.
spectra. In particular, the monitoring of meth-
ane emissions in the near-infrared range The relative abundances of the newly formed
showed a decrease of the temperature from species are indicated in Table 1.
1,000 K at t = 10 min to 600 K at
t = 25 min. HST images showed the forma- Long-Term Evolution of Impact Phenomena
tion of dark craters surrounded by crescent- The new molecules formed by shock chemistry
shaped areas resulting from infalling ejecta, had very contrasting lifetimes. Water was the
indicating the formation of dust, possibly first molecule to disappear, after a few hours.
including carbonaceous and silicate material Still, the water from the SL9 collision is
(Fig. 2). believed to be at least partly responsible for the
traces of stratospheric water discovered in 1997
New Molecules by the Infrared Space Observatory and later
Several species new to Jupiter have been detected observed by other submillimeter satellites.
in the stratosphere during the splash phase: OCS, NH3, and S2 were detected during a few
weeks. CO and CS2 were observed during sev-
• The most abundant was CO, detected both in eral months, and HCS and CN were detectable
the infrared and in the millimeter range. In over several years. The observed lifetimes were
particular, infrared spectra of CO showed a found to be in good overall agreement with
high excitation temperature of a few thousand the predictions inferred from photochemical
K at the beginning of the splash phase. models.
• Water was detected in the near infrared by the The dust observed in the impact craters could
(NIMS) Near Infrared Mapping Spectrometer be monitored during the weeks and months fol-
instrument aboard Galileo and by the Kuiper lowing the collision. The clouds first extended in
Airborne Observatory, with an excitation tem- longitude as an effect of Jupiter’s fast rotation
perature of 1,000 K. and, after a couple of weeks, formed a continuous
• Sulfur species were identified by the UV spec- band at latitude 44S. During about a year, the
trometer of the HST: S2, CS2, CS, and possibly clouds extended in latitude between 20S and 80-
H2S. S. Aerosols, first formed in the stratosphere,
Comet Shoemaker-Levy 9 523
Comet Shoemaker-
Levy 9, Fig. 2 One of the
first impact sites of the SL9
collision as observed by the
HST. All impacts occurred
at latitude 44S.
The enlargement on the left
side shows the crescent- C
shaped structure
surrounding the main
impact crater in the
atmosphere (# NASA)
Comet Shoemaker-Levy 9, Table 1 New molecules formed in Jupiter atmosphere by shock chemistry
Molecular mass in SL9 Elemental mass in Elemental mass in a 1015 g
Molecule Observations impact (g) SL9 impact fragment (comet)
H2O IR, (radio) >2 1012 >1.5 1014 [O] 5 1014 [O]
CO (IR), radio 2.5 1014
NH3 (UV), IR 1 1013 1 1013 [N] 2 1013 [N]
HCN IR, radio 6 1011
S2 UV 1.5 1012
CS2 UV 1.5 1011
CS (UV), radio 5 1011 5 1012 [S] 4 1013 [S]
H2S UV Marginal
OCS Radio 3 1012 1 1014 [C] 2 1014 [C]
Silicate IR 6 1012 8 1013 [Si] 1 1014 [Si]
4 1013 [Mg] 4 1013 [Mg]
moved downward, contributing to the atmo- cross the radiation belts, intercepting the disk of
spheric profile cooling, to reach tropospheric Jupiter around latitude 44S.
levels after about a year. Auroral phenomena were also recorded, in
some cases, prior to the impacts, implying a strong
Magnetospheric Effects heating of the upper stratosphere, in particular UV
The Jovian ▶ magnetosphere was also affected emissions of H and H2, and H+3 in the infrared.
by the collision. The synchrotron radiation, mon- Often, auroral emissions were detected at the loca-
itored at centimeter wavelengths, showed an tion of the northern counterparts of the impact
enhancement of a few tens of percent during the sites, corresponding to the footprints of the
week of impacts. The emission was mostly connecting magnetic field lines: X-ray emissions
observed at localized longitudes, between were recorded by the ROSAT satellite at the time
100 and 240 , showing evidence for a new pop- of two impacts; UV emissions were observed by
ulation of excited electrons. This region corre- the HST; and H+3 near-IR emissions were detected
lates with the side where the magnetic field lines from the terrestrial ground-based facilities.
524 Comet Shower
About the SL9 Comet 10 mm is also in agreement with typical cometary

All astrometric measurements performed before values. In conclusion, several facts support a
and during the collision were used to retrieve the cometary rather than asteroidal origin for the
best orbital fits ever obtained for a comet. The impactor: the small size, the weak but real level
progenitor of comet SL9 had been captured by of activity, the low density, the very low tensile
Jupiter nearly in the year 1930. Before its capture, strength, the chemical composition, and the pres-
the comet’s orbit was probably within Jupiter’s ence of silicates. Comet SL9 was actually a very
orbit, with a low eccentricity and a low small and very common Jupiter-family comet.
inclination. In summary, the collision of comet SL9 with
From the disruption of the comet’s nucleus in Jupiter has not allowed astronomers to fully
1992, its tensile strength was inferred. The very understand the nature and the composition of
low value, 100 Pa, is consistent with a fluffy the comet; however, it provided a real-time
aggregate of submicron particles. Models of the observation of the response of a planetary atmo-
swarm elongation are consistent with a progenitor sphere to a large meteoritic impact.
of 1.5 km in size, with a density of 0.5 g/cm3.
These numbers are consistent with the estimates
inferred for the impactors on the basis of the See Also
impact dynamical models. The comet’s activity
was measured in 1993 and 1994 before the impact. ▶ Comet
Very low dust production rates were estimated ▶ Jupiter
(1–5 kg/s for the different impacts). No gaseous
activity was detected; an upper limit of 1027 mol/s
was retrieved for the OH production rate. References and Further Reading
Information about the comet’s composition
was obtained from visible spectra taken during Encrenaz T (2001) Shoemaker-Levy-Jupiter collision. In:
some impacts that showed a variety of atomic Murdin P (ed) Encyclopedia of astronomy and astro-
physics. IoP Publishing, Bristol, pp 2413–2420
lines (Fe, K, Ca, H, Na, Mg, Mn, Cr, and for the
Noll KS, Weaver HA, Feldman PD (eds) (1996) The col-
first time, Li). UV spectra taken by the HST also lision of comet Shoemaker-Levy and Jupiter. Cam-
showed atomic and ionic transitions: H, He, S, Si, bridge University Press, Cambridge
Mg+, Fe+, Si+, and Al+. All these species, absent Spencer KR, Mitton J (eds) (1995) The great comet crash.
from Jupiter’s spectrum, belong to the impactor;
West RM, Bohnhardt H (eds) (1995) Proc European work-
they are frequently observed in the spectra of shop on the SL9-Jupiter Collision, no. 52, Garching,
sun-grazing ▶ comets. ESO
The composition of SL9 is difficult to retrieve
from the new molecules formed during the
impacts because they result from a recombination
of products partly possibly having a planetary Comet Shower
origin. Still, the oxygen and sulfur-bearing mol-
ecules, absent from Jupiter’s stratosphere, can be Sean N. Raymond
used to infer the O and S content of the impactor. Laboratoire d’Astrophysique de Bordeaux,
The inferred O/S is consistent with cometary CNRS, Universite de Bordeaux, France
values. The carbon content of the impactor can-
not be inferred as the new molecules may have
been formed from Jovian methane. Assuming Definition
that all the nitrogen of the new molecules is
coming from the impactor, the inferred N/O A comet shower is a short-lived burst in the flux
ratio is consistent with a cometary origin. The of comets entering the inner Solar System from
Si/O ratio inferred from the silicate signature at the ▶ Oort cloud. Comet showers are thought to
Comets, History of 525
be triggered primarily by relatively close pas-

sages of nearby stars with the Solar System. Comet Wild 2
Comet showers are not considered an impact
threat on Earth. Therese Encrenaz
See Also C
▶ Oort Cloud Definition
▶ Comet 81 P/Wild, also named Wild 2, was

discovered in 1978. After a passage close to Jupi-
ter in 1974, its orbit moved toward the inner solar
Comet Tempel 1 system, with a high ellipticity and a short period
(6.4 years). Its perihelion is close to 1 AU. In
Therese Encrenaz January 2004, comet Wild 2 was approached by
LESIA, Observatoire de Paris - Section de the Stardust spacecraft which collected cometary
Meudon, Meudon, France grains from its tail and brought them back to
Earth in 2006. This cometary matter included
material originating from different regions of
Definition the solar system before being incorporated into
the comet nucleus. The presence of glycine, the
Comet 9/P Tempel 1 was discovered in 1867 at simplest amino acid, has been reported in the
the Observatoire de Marseille by the astronomer cometary samples. Comet Wild 2 is the only
Ernst Wilhelm Tempel. Its orbit, very close to the comet for which samples have been brought
▶ ecliptic, has a period of 5.5 years around the back to Earth and analyzed.
Sun and lies between the orbits of Mars and
Jupiter. The diameter of the comet is 6.5 km and
its rotation period is 41 h. Comet Tempel 1 has
been explored in detail by the NASA probe See Also
▶ Deep Impact which sent an impactor to its
surface on July 4, 2005. The impact led to the ▶ Comet
formation of a 30-m large crater and the massive ▶ Stardust Mission
ejection of dust, water, and HCN. Following the
success of another cometary mission, the ▶ Star-
dust encounter of comet Wild 2 in 2004, NASA
decided in 2007 to reorient the Deep Impact
spacecraft for a new flyby of comet Tempel Comets, History of
1. The flyby successfully took place on February
15, 2011, with a closest approach of less than Stéphane Le Gars
200 km, and the spacecraft recorded several tens Centre François Viète, Université de Nantes,
of images of the surface, including the impact Nantes, France
crater site.
Keywords
See Also
Astronomy; Comets; Gravitation; Halley;
▶ Comet History; Meteorites; Oort; Physics
526 Comets, History of
History (typically photodissociation products). It is only

in 1950 that the American astronomer Fred
Since ancient times, ▶ comets have raised ques- Whipple described the comets as “dirty snow-
tions and polemics. Visual observations of balls,” speculating that their nucleus is essen-
comets date back to the eleventh century BCE, tially constituted of watered ice mixed with
when Chinese chronicles related a cometary carbonic ice, silicates, and organic compounds.
apparition to war between two kings. There are It is also in 1950 that Dutch astronomer Jan Oort
many references in classical literature, including calculated that long-period comets came from a
Pliny’s report of a comet during the Battle of gigantic spherical reservoir, containing thou-
Salamis (480 BCE). During the Middle Ages in sands of millions of those little bodies, and
Europe, comets were seen as portents, of the located them at 1 or 2 light years from the Sun.
Norman conquest of England in 1066 in the Starting with the return of Comet Halley in 1986,
Bayeux Tapestry and of the birth of Jesus as radio and infrared astronomers have identified
depicted by Giotto in 1304. The New World many molecular constituents of comets by
also linked comets and important events, includ- observing their emission at millimeter wave-
ing that observed by the Aztec emperor Mocte- lengths. In 2006, the Stardust space probe
zuma, prior to the conquest of Mexico by Cortes. brought back to Earth dust from the Wild-2
If the Chaldean considered them as planets, some comet, out of which analysis led to new knowl-
Greek philosophers such as Aristotle situated edge on the solar system.
them in the area between the Earth and the
Moon. From the sixteenth century, comets were
being studied more systematically. Their move- See Also
ments were described through the constellations;
an interest is taken in their tail’s orientation with ▶ Comets, History of
regard to the Sun, and an estimation of their ▶ Kuiper Belt
parallax puts them definitely beyond the Moon ▶ Oort Cloud
(the turning point in the study of comets was
made with Tycho Brahe’s observations of the
parallax of the comet of 1577). References and Further Reading
It is since the universal gravitation law, pro-
posed by Newton in 1687 in his Philosophiae Acker A (2005) Astronomie astrophysique. Introduction.
Dunod, Paris
Naturalis Principia Mathematica, that the
Bosler J (1928) Cours d’Astronomie. III Astrophysique.
astronomer Edmund Halley identified the 1682 Librairie scientifique Hermann et Cie, Paris
comet with those of 1607 and 1531 and predicted Durán D Historia de las Indias de Nueva España y Islas de
its return for 1759: at this moment, it was proved Tierra Firme, Fondos de la Biblioteca Nacional
(Spain)
that those celestial bodies were bound, like Flammarion C (1880) Astronomie populaire. C. Marpon et
planets, by gravitation, describing on the other E. Flammarion, Paris
hand elliptical (in that case they are periodical), Henarejos P (1999) Comètes. In: Dictionnaire de
parabolic, or hyperbolic orbits about the Sun. l’Astronomie, Encyclopedia Universalis et Albin
Michel, Paris, p 164–175
Then, in the nineteenth century, attention turned
Millochau G (1910) De la Terre aux astres. Librairie
to the origin and the composition of those heav- Delagrave, Paris
enly bodies. Analysis of the spectrum and polar- Olson R, Pasachoff J (1987) New information on comet
ization of light permitted astronomers at this P/Halley as depicted by Giotto di Bondone and other
Western artists. Astron Astrophys 187:1–11
time to identify in the comet’s coma
Yeomans D (1991) Comets: a chronological history
(atmosphere; from the Latin word for “hair”) of observation, science, myth, and folklore. Wiley,
and tail both dust and atoms and small molecules New York
Compatible Solute 527
Common Ancestor Community Genome
Luis Delaye ▶ Metagenome

Departamento de Ingenierı́a Genética,
CINVESTAV-Irapuato, Irapuato, Gto, Mexico
C
Synonyms
Compatible Solute
Josefa Anton
Concestor; Most recent common ancestor;
Department of Physiology, Genetics and
MRCA
Microbiology, University of Alicante, Alicante,
Spain
Definition
Keywords
A common ancestor is the ancestral biological
entity from which a group of other biological
Haloadaptation; Halophile; Halotolerant
entities have evolved. Examples of biological
entities are species, biological structures, or
molecular ▶ sequences. One of the main ideas
Synonyms
of Charles Darwin in The Origin of Species
(1872) is that species are related by common
Osmolyte
ancestry, i.e., that species have evolved by diver-
gence from a common ancestral specie. As a
result of evolution by divergence from a common
Definition
ancestor, the phylogenetic relationships among
species or other biological entities can be
A compatible solute is a substance compatible
described using a treelike structure. The related
with the cellular metabolism that accumulates in
concept of most recent common ancestor
the cytoplasm to balance external osmotic pres-
(MRCA) specifies the least ancient common
sure. This accumulation can be due either to
ancestor.
transport from the medium or to de novo synthe-
sis and helps maintaining turgor pressure, cell
volume, and concentration of electrolytes, all
See Also needed for cell viability and proliferation.
▶ Cenancestor
▶ Darwin’s Conception of the Origins of Life Overview
▶ Endosymbiosis
▶ Evolution, Biological Microorganisms cope with osmotic stress (due
▶ Homology to high salt, freezing, and/or desiccation) by
▶ Last Universal Common Ancestor using two different types of strategies. Archaea
▶ Phylogenetic Tree of the order Halobacteriales, anaerobic Bacteria
▶ Phylogeny belonging to the order Haloanaerobiales,
▶ Sequence and some members of the Bacteroidetes
528 Complex Organic Molecules
(i.e., Salinibacter ruber) accumulate high con- References and Further Reading
centrations of inorganic ions (mostly potassium)
in the cytoplasm. This is known as the “salt-in” Empadinhas N, da Costa MS (2008) Osmoadaptation
mechanisms in prokaryotes: distribution of compatible
strategy and requires the adaptation of the entire
solutes. Int Microbiol 11:151–161
intracellular machinery in order to function Roberts MF (2005) Organic compatible solutes of
in this highly saline environment. The second halotolerant and halophilic microorganisms. Saline
strategy relies on intracellular accumulation, Syst 1:5
either by uptake or de novo synthesis, of
low-molecular-weight organic compatible sol-
utes (also called osmolytes) to balance the exter- Complex Organic Molecules
nal osmotic pressure. The accumulation of such
solutes is very widespread in nature since the Kensei Kobayashi
cytoplasm of most organisms does not tolerate Yokohama National University, Tokiwadai,
salt. In addition, this is a very versatile strategy Hodogayaku, Yokohama, Japan
that allows for rapid response to changing envi-
ronments since organisms can regulate their
intracellular concentration according to the sur- Definition
rounding salinity.
Compatible solutes are small organic mole- In astronomy, complex organic molecules are
cules that act as osmoprotectants, thanks to their defined as molecules with several carbon atoms
ability to stabilize cellular proteins, providing a such as benzene or acetic acid. These molecules
hydration shell and stabilizing their tertiary struc- have been detected in interstellar space using
tures without interfering in cell metabolism (this radio telescopes. A gas mixture of methane and
is why they are called “compatible”). Indeed, nitrogen, for example, yields complex organic
compatible solutes not only provide protection compounds upon UV irradiation or exposure to
from osmotic stress but some can also act as an electric discharge. These are sometimes
thermostabilizers, a property that has been referred to as ▶ tholins, and they can be formed
exploited for biotechnological purposes. abiotically in the atmosphere of ▶ Titan (the
There is a wide range of compatible solutes, largest satellite of Saturn).
but they can be classified into a few chemical In chemistry, “complex organic molecules”
categories (Empadinhas and da Costa 2008) generally refer to much larger polymer-like mol-
such as amino acids and derivatives (including ecules such as ▶ proteins. Proteins are typical
ectoines and hydroxyectoines), sugars and deriv- complex organic polymers, with well-defined
atives, phosphodiesters, and polyols. Some of three-dimensional structures, composed of 20 dif-
them, such as glycine betaine, are universal com- ferent amino acids. Given the immense possible
patible solutes, while others are restricted to a variety of these polymers, they are indeed
specific group of microorganisms. For a detailed “complex.”
description of the chemical nature and occur-
rence of organic compatible solutes within halo-
philic and halotolerant organisms, see See Also
Roberts (2005).
▶ HCN Polymer
▶ Insoluble Organic Matter
See Also ▶ Protein
▶ Radio Astronomy
▶ Halophile ▶ Tholins
▶ Halotolerance ▶ Titan
Complexity 529
Overview
Complex Organic Product
The study of complexity and complex systems is
▶ Tholins so broad and encompasses so many disciplines
that it is difficult to define. There are different
definitions suitable for different contexts and pur-
poses. Etymologically, complexity comes from C
the Latin plexus, which means interwoven.
Complex Organisms A complex system is one in which elements
interact in such a way that it is difficult to separate
▶ Multicellular Organisms their behavior. In other words, if one element
affects the state of another element, the dynamics
of the system cannot be reduced to the states of
the elements, since interactions are relevant for
the future state of the system. Examples of com-
Complexity plex systems include a ▶ cell, a brain, a city, the
Internet, a market, a crowd, an ▶ ecosystem, a
Carlos Gershenson ▶ biosphere, and an atmosphere. A cell is com-
Instituto de Investigaciones en Matemáticas posed of molecules, but the behavior of a cell
Aplicadas y en Sistemas, Universidad Nacional cannot be reduced to that of molecules. Their
Autónoma de México, DF, Mexico interactions generate constraints and information
that is not present in molecules and determine the
behavior of the cell.
Keywords Some approaches to complexity do not focus
on its systemic aspect, but more on its probabi-
Complex systems; Computation; Information; listic or algorithmic aspect. These are related with
Interactions; Nonreductionism; Scientific information theory, e.g., how probable is a string
paradigm of bits, how long is the shortest algorithm that
produces a string of bits, what is the shortest time
it can take an algorithm to produce a string of bits,
Definition or how compressible is a string of bits. Intui-
tively, in most of these descriptions, complexity
There is no single definition of complexity represents a balance between order (stability) and
(Edmonds 1999; Gershenson 2008; Mitchell chaos (variability) (Kauffman 1993).
2009), as it acquires different meanings in differ- Since complexity can be found in almost any
ent contexts. A general notion is the amount of field, some people question its usefulness, while
information required to describe a phenomenon others defend it as a novel scientific paradigm
(Prokopenko et al. 2009) (note that this depends that complements the traditional reductionist
on the scale (Bar-Yam 2004) and context in approach (Gershenson and Heylighen 2005;
which the description is made, e.g., an organism Morin 2006).
requires more information to be described at a
molecular scale than at a population scale), but it
can also be understood as the length of the Basic Methodology
shortest program required to compute that
description, as the time required to compute that There have been many methods developed within
description, as the minimal model to statistically the study of complexity that have proven to be
describe a phenomenon, etc. very useful, since they are able to take into
530 Complexity
account the interactions of the elements of a Applications

complex system. Tools include agent-based
modeling, networks, ▶ cellular automata, The scientific study of complexity and complex
▶ genetic algorithms, and swarm intelligence. systems has found applications in physical,
Most of complexity research is based on com- chemical, biological, computational/informa-
puter simulations. On the one hand, complex tional, social, economic, engineering, and other
models tend to involve large numbers of elements fields. In many cases, the concepts, tools, and
and/or interactions, which are difficult to handle methods of complexity have been applied to spe-
without computer aid. On the other hand, inter- cific problems, e.g., self-assembly, pattern forma-
actions generate novel and relevant information tion, adaptive control, protein folding, ecological
that is not present in initial or boundary condi- studies, robotics, evolution, etc. In other cases,
tions. This makes it difficult to know a priori a the study of complexity per se has also attracted
final state of a system without computing all of its broad attention.
transitions, i.e., predictability is limited. A model Complexity formalisms allow the study of
has to “run” before something definitive can be phenomena at different scales and to relate them
said about it. Cellular automata provide a clear under the same framework. This is useful when
example of this. Thus, an equation-based multiple scales (spatial, temporal, functional,
approach is in many cases insufficient to explore dynamical) interact within a system, since the
the properties of a model. same language can be used to relate the scales.
There are many concepts that are related to the This is not feasible with a reductionist approach.
study of complexity, such as nonlinearity, self-
organization, adaptation, chaos, and emergence.
Future Directions
Key Research Findings Some have speculated that complexity is a fad

and it will lose its popularity, following the steps
The scientific study of complexity has increased of similar movements: cybernetics, catastrophe
the understanding of phenomena in many differ- theory, and chaos theory. Nevertheless, complex-
ent fields. Common examples include models of ity has been studied (under this name) since the
collective behavior, complex networks 1980s, and everything indicates that the interest
(molecular, metabolic, genetic, neural, trophic, in it is growing. The concepts and methods that
ecologic, social, economic, organizational, polit- have been developed within the study of com-
ical, geographical), nonlinear dynamics, evolu- plexity and complex systems are permeating into
tion, and distributed systems. Theoretically, all disciplines. Maybe people will not use the
complexity has also provided several concepts, term complexity, but this is not relevant. Com-
formalisms, and tools. plexity is helping shape a shift in the scientific
The main difference of complexity-related worldview, from reductionist to “interactionist.”
and traditional techniques is that complexity can This is relevant, since this shift is allowing us to
easily include millions of variables into consid- expand the frontiers of our knowledge.
eration, e.g., with cellular automata, multi-agent
systems, or networks. This is difficult to achieve
with, e.g., differential equations, which are more See Also
suitable for contexts where there are few vari-
ables considered and the state space or phase ▶ Artificial Life
space does not change, i.e., is stationary. The ▶ Cellular Automata
tools of complexity are suitable for studying ▶ Emergence of Life
nonstationary spaces, i.e., those that change ▶ Genetic Algorithms
with time. ▶ Scale-Free Networks
Composomes 531
References and Further Reading Composomes arise in an artificial chemistry

description of early evolution, prior to the emer-
Bar-Yam Y (1997) Dynamics of complex systems. Stud- gence of complex polymeric molecules such as
ies in nonlinearity. Westview, Boulder
RNA or proteins. This is a stage at which random
Bar-Yam Y (2004) Multiscale variety in complex systems.
Complexity 9(4):37–45 assemblies of simple molecules might have
Edmonds B (1999) Syntactic measures of complexity. acquired a prebiotic metabolism before exact
Doctoral thesis, University of Manchester, Manchester polymeric replication existed. The dynamics of C
Gershenson C (ed) (2008) Complexity: 5 questions. Auto-
such molecular assemblies are thought to have
matic Press/VIP, Copenhagen. ISBN 8792130135
Gershenson C, Heylighen F (2005) How can we think the been governed by processes such as catalytic
complex? In: Richardson K (ed) Managing organiza- closure, affording the emergence of mutually cat-
tional complexity: philosophy, theory and application. alytic sets (Kauffman 1993) or a transition from a
Information Age, Greenwich, Chapter 3
disordered to an ordered quasistationary state
Holland JH (1995) Hidden order: how adaptation builds
complexity. Helix Books/Addison-Wesley, Reading (Dyson 1999), both mimicking self-replication.
Kauffman SA (1993) The origins of order. Oxford Uni-
versity Press, Oxford
Meyers RA (ed) (2009) Encyclopedia of complexity and
systems science. Springer, New York
Overview
Mitchell M (2009) Complexity: a guided tour. Oxford
University Press, Oxford The specific computer-simulated model within
Morin E (2006) Restricted complexity, general complex- which composomes have been invoked is the
ity. In: Gershenson C, Aerts D, Edmonds B (eds) Phi-
Graded Autocatalytic Replication Domain
losophy and complexity, worldviews, science and
us. World Scientific, Singapore (GARD) model (Segré et al. 2000; Markovitch
Prokopenko M, Boschetti F, Ryan A (2009) An and Lancet 2014). This model simulates assem-
information-theoretic primer on complexity, self- blies of small molecules, typically lipids and
organisation and emergence. Complexity 15(1):11–28
other amphiphiles, that spontaneously accrete
into discrete assemblies, much in line with
Oparin’s Conception of Origins of Life in a pri-
mordial soup environment. GARD is intimately
Composomes linked to the concept of a Lipid World (Segré
et al. 2001). This is an origin of life scenario
Doron Lancet that focuses on amphiphilic micelles and vesi-
Weizmann Institute of Science, Rehovot, Israel cles, in line with lipid protocells and other types
of prebiotic protocells.
A key concept in GARD is that in a primordial
Keywords soup, a huge diversity of amphiphiles has been
available, so molecular assemblies could assume
Prebiotic metabolism; Self-replication; Mutually a large variety of compositions. The GARD
catalytic sets; Selection; Evolution; Protocells; chemical kinetics formalism shows that very spe-
Vesicles cific compositions may breed true (with muta-
tions), in a process of repeated accretion-driven
growth followed by physical fission. Such
Definition favored compositions are defined as
composomes, a name hinting at the idea that
A composome is a quasistationary state in a assemblies are capable of genome-like composi-
dynamic description of compositional molecular tional information transmission from one gener-
assemblies (Segre et al. 2000). The term origi- ation to another. Composomes are a cornerstone
nated in the context of origin of life, in a of more recent studies that portray, among others,
monomer-world scenario (Shapiro 2007), pro- a route from metabolism-like composomes to
posed as an alternative to the RNA world. genetic replication (Norris et al. 2012) and early
532 Computational Biology
ecology (Hunding et al. 2006; Markovitch and Segré D et al (2001) The lipid world. Orig Life
Lancet 2014). While some question the capacity Evol Biosph 31:119–145
Shapiro R (2007) A simpler origin for life. Sci Am 296:46
of mutual catalysis-driven compositional assem- Vasas V et al (2010) Lack of evolvability in self-
blies devoid of informational biopolymer, to sustaining autocatalytic networks constraints
undergo selection and evolution (Vasas et al. metabolism-first scenarios for the origin of life. Proc
2010), solid evidence for the existence of such Natl Acad Sci 107:1470–1475
capacities has been reported (Markovitch and
Lancet 2012).
The origin of life is ridden with a chicken or Computational Biology
egg problem: present-day RNA and DNA are
rather inert in the absence of catalytic proteins, ▶ Bioinformatics
and proteins cannot be formed without
biopolymeric guidance. Composomes may be a
clue as to where to look for a solution to this
riddle (Shapiro 2007). CONAE, Argentina
Michel Viso
See Also CNES/DSP/SME, Vétérinaire/DVM, Astro/
Exobiology, Paris Cedex 1, France
▶ Artificial Chemistries
▶ Chicken or Egg Problem
▶ Lipid Synonyms
▶ Metabolism, Prebiotic
▶ Oparin’s Conception of Origins of Life Argentina space agency
▶ Origin of Life
▶ Primordial Soup
▶ Protocell Definition
▶ RNA World
The “Comisión Nacional de Actividades
References and Further Reading Espaciales” (CONAE) is the space agency for
Argentina. Argentina’s first activities in the
Dyson F (1999) Origins of life. Cambridge University space field date back to 1961, when the National
Press, Cambridge Commission for Space Research (Comisión
Hunding A et al (2006) Compositional complementarity
and prebiotic ecology in the origin of life. Bioessays
Nacional de Investigaciones Espaciales, CNIE)
28:399–412 was first established within the Argentine air
Kauffman SA (1993) The origins of order: self- forces area. With other local and international
organization and selection in evolution. Oxford Uni- organizations, CNIE carried out, by means of
versity Press, New York
rockets and stratospheric balloons, the first
Markovitch O, Lancet D (2012) Excess mutual catalysis is
required for effective evolvability. Artif Life Southern Hemisphere scientific atmospheric
18:243–266 studies, which included wind measuring and
Markovitch O, Lancet D (2014) Multispecies population assessment of neutral atmosphere dynamics
dynamics of prebiotic compositional assemblies.
J Theor Biol 357:26–34
using the alkaline clouds technique. Together
Norris V et al (2012) How did metabolism and genetic with the Argentine Institute of Aeronautics and
replication get married? Orig Life Evol Biosph Space Research, CNIE designed and constructed
42:487–495 a family of one- and two-stage sounding rockets.
Segré D et al (2000) Compositional genomes: prebiotic
information transfer in mutually catalytic noncovalent
In 1991, the Argentine Government decreed the
assemblies. Proc Natl Acad Sci U S A creation of this National Commission for Space
97(8):4112–4117 Activities as a civil organization. Since 1996, this
Concretions (Mars) 533
specialized agency accomplishes its mission membranes like those of neurons. Hydrogen ion
governed by the Ministry of Foreign Affairs. gradients are generated by the electron transport
CONAE is involved, in particular, in use of systems embedded in membranes, either as a pH
space data for telemedicine and tele- gradient or a ▶ membrane potential. For instance,
epidemiology and use of space infrastructure for a gradient equivalent to 210 mv or 3.5 pH units
health emergencies during disaster management. (1,000-fold gradient) is the source of energy
driving ▶ ATP synthesis in mitochondria, chlo- C
roplasts, and bacterial membranes. Although
concentration gradients are essential to life
Concentration Gradients today, it is uncertain what role they may have
played in the origin of life. Early membranes
David Deamer were likely to be relatively permeable to solute
Department of Chemistry, University of ions, so that primitive cells would be unable to
California, Santa Cruz, Santa Cruz, CA, USA use gradients as an energy source.
Keywords See Also
Membrane potential; Semipermeable membrane; ▶ Membrane Potential

Stored energy
Definition Concestor
Concentration is a measure of the amount of ▶ Common Ancestor

solute in a solvent, typically expressed in units
of moles per liter. A 1.0 molar solution (abbrevi-
ated 1.0 M) contains 1 mole of a solute in 1 liter of
total volume. A concentration gradient exists Concretions (Mars)
when a higher concentration of a solute is sepa-
rated from a lower concentration, by a semiper- Alessandro Airo
meable membrane. Institut f€ur Geologische Wissenschaften
Tektonik und Sedimentäre Geologie, Freie
Overview Geowissenschaften, Berlin, Germany
Concentration gradients of solutes are common in

living cells and are essential sources of energy for Definition
all forms of life. Concentration gradients are gen-
erated and maintained across biological mem- Concretions are postdepositionally cemented
branes by ion pump enzymes that transport ionic areas within sedimentary rocks that occur as
solutes such as sodium, potassium, hydrogen spheres, disks, tubes, botryoidal aggregates, or
ions, and calcium across the membrane. Energy have an irregular lumpy shape. The cement
is required to produce a gradient, so the gradient forms through precipitation from pore water and
is a form of stored energy. An important example is usually composed of carbonate (e.g., calcite,
is the sodium and potassium ion gradient across siderite), sulphate (e.g., gypsum), silica (e.g.,
most cell membranes, which produces the resting chert), or iron oxide (e.g., hematite). Nodules
potential and action potentials of excitable have a similar origin, but instead of cementing
534 Condensate Layer
the pre-existing sediment in place, they substitute Definition

the particles through relocation or dissolution.
The Mars Exploration Rover Opportunity was In planetary science, the condensation sequence
the first to discover concretions on Mars, the refers to the order in which chemical compounds
so-called ‘blueberries’, which are cemented by transition from gas to solid phase in a protoplan-
hematite (Squyres et al. 2004). The Mars Science etary nebula (disk), based on the condensation
Laboratory Curiosity has discovered light-toned temperature of each compound. The condensa-
concretions within mudstones at Gale crater, tion sequence is important because the tempera-
which possibly are cemented by sulphates ture of a protoplanetary nebula varies radially and
(Grotzinger et al. 2014). The occurrence of con- temporally, and the chemical composition and
cretions on Mars is insofar of interest to astrobi- mass of any bodies formed at a given place and
ologists as they indicate the former presence of time is defined by the materials that can condense
groundwater. at the local temperature.
See Also
Overview
▶ Aquifer (Mars)
▶ Meridiani (Mars) Condensation is defined as the process by which
chemical materials change phase from the gas-
eous or liquid phase to the solid phase. Chemical
References
condensation occurs within a narrow range of
Grotzinger JP et al (2014) A habitable fluvio-lacustrine temperature specific to each individual chemical
environment at Yellowknife Bay, gale crater, mars. compound (a few tens of degrees), defined by the
Science 343:386–387 abundance of the elements in the nebular gas and
Squyres SW et al (2004) The opportunity Rover’s Athena the molecular structure and composition of the
science investigation at meridiani planum, mars. Sci-
ence 306:1698–1703 solid compound. Effective condensation temper-
atures depend on the local environment (i.e.,
pressure) and can be lowered by including cata-
lytic reactions (i.e., grain growth reactions).
Condensate Layer A “condensation sequence” orders a specific list
of chemical compounds of interest by their con-
▶ Clouds densation temperatures, in order to determine
which materials will condense in a medium of a
specific temperature.
The Solar System is thought to have formed
Condensation Sequence from a rotating disk of gas and dust called the
“▶ solar nebula” (this is true for other planetary
Avi M. Mandell systems as well; we call these disks “protoplane-
NASA Goddard Space Flight Center, Greenbelt, tary disks”). The temperature of the solar nebula
MD, USA varied both radially (from the inner regions of the
disk to the outer regions) and vertically (from the
midplane to the surface) due to heating from the
Keywords Sun and viscous heating at the midplane. The
temperature in different parts of the solar nebula
Cosmochemistry; Solar nebula also changed over time as the disk material was
Conjugation 535
depleted due to accretion to the star and

photoevaporation. The material condensing out Condensation Temperature
of the gaseous solar nebula into grains and larger
bodies would therefore be a function of both William M. Irvine
location and of age, resulting in a changing chem- University of Massachusetts, Amherst, MA, USA
ical composition of bodies over space and time in
the nebula. A typical condensation sequence of C
the nebular gas starts with refractory oxides and Definition
silicates (1,700–1,400 K), followed by iron and
nickel which will eventually segregate into plan- The condensation temperature is that at which a
etary cores and the silicates of their future mantle given gas-phase constituent condenses into a liq-
and crust (1,350–1,150 K). Alkali elements and uid. This temperature depends on the physical
sulfide-loving (chalcophile) elements such as Cu, and chemical state of the system.
Zn, and Pb and finally volatile elements such as S,
C, N, and H2O join between 600 and 100 K. The
temperature at which condensation came to a halt
for the material of a planetary body, therefore,
defines its depletion in volatiles, among them Conjugation
water.
One of the most interesting consequences of Carlos Briones
the condensation sequence is therefore the pro- Centro de Astrobiologı́a (CSIC/INTA), Consejo
posed existence of a “▶ snow line” in the solar Superior de Investigaciones Cientı́ficas, Madrid,
nebula. The snow line is defined as the radial Spain
location in the solar nebula beyond which the
local temperature of the nebula drops below the
condensation temperature of water. Bodies Definition
formed beyond this boundary would contain
water, while bodies inside this boundary would In microbiology, conjugation is the process by
be relatively waterpoor. In fact, Earth is thought which two individuals of the same – or even
to have formed inside the snow line, hence, the different – species exchange their genetic mate-
concept of “water delivery”. The existence of rial during a temporary union. Among ▶ bacte-
water ice beyond the snow line has increased ria, conjugation is a widespread mechanism that
the solid surface density at these locations and allows the transmission of one conjugative
led to the rapid growth of the giant planets’ ▶ plasmid from one cell, called “donor,” to
cores. another, the “recipient,” through the physical
contact between them. Conjugative plasmids
carry genes that promote bacterial conjugation
and are frequently involved in ▶ lateral gene
See Also transfer – also called horizontal gene transfer,
HGT – between individuals of different species.
▶ Core Accretion, Model for Giant Planet For example, antibiotic resistance or the ability
Formation to metabolize a new organic molecule can be
▶ Protoplanetary Disk transmitted by conjugation among microorgan-
▶ Snow Line isms. In the majority of bacteria, conjugation is
▶ Solar Nebula not a standard mode of reproduction as it is the
▶ Water, Delivery to Earth case in sexual organisms. In turn, in certain
536 Constructive Biology
▶ protists such as paramecia, although they usu-

ally reproduce asexually by fission, conjugation Continental Crust
is a sexual process of reproduction between
individuals of opposite mating types. Conjuga- Nicholas Arndt
tion is also used as a genetic engineering tech- ISTerre, Université Grenoble Alpes, France
nique in the lab.
Definition
See Also
Continental crust is that portion of the Earth’s
▶ crust composed mainly of low-density siliceous
▶ Bacteria
(granitoid) rock. It occupies about one-third of the
▶ Domain (Taxonomy)
total crust and underlies most of dry land. Its thick-
▶ Genetics
ness varies from
10 km in rifts to up to 80 km
▶ Lateral Gene Transfer
beneath mountain belts. Dominant constituents are
▶ Plasmid
granitic rocks and their metamorphic equivalents
▶ Protists
(gneiss) and metasedimentary or metavolcanic
▶ Transformation
sequences. Continental crust forms above ▶ sub-
duction zones and contains a large proportion of
the Earth’s budget of incompatible trace elements
and heat-producing elements. The presence of con-
tinental crust is a signature of ▶ plate tectonics that
Constructive Biology
distinguishes ▶ Earth from other planets.
Because continental crust is made of lighter
▶ Synthetic Biology
(granitoid) material than (basaltic) oceanic crust
and its overall density is therefore less, the litho-
spheric segments that contain continental crust
float higher on the ▶ asthenosphere than those
without it. Liquid water on Earth thus usually
Contamination, Probability
only covers its margins (“continental margins,”
shelves), allowing there the colonization of
Catharine A. Conley
regions covered by shallow water and within the
photic zone. This gives rise to the “carbonate
factory” on continental shelves.
Definition
See Also
For purposes of ▶ planetary protection, the prob-
ability of contamination is determined, using ▶ Crust
a formulation of the ▶ Coleman-Sagan equation ▶ Earth
that an Earth-originating organism might grow ▶ Granite
and propagate in another planetary environment. ▶ Plate Tectonics
See Also Continental Lithosphere
▶ Coleman-Sagan Equation ▶ Continents

Continents 537
trace elements, including the sources of radioac-

Continental Plate tive heat. Where continents collide with other
plates, their edges thicken to form mountain
▶ Continents belts, which eventually erode to fill sedimentary
basins.
C
Overview
Continental Tectosphere
From a geological perspective, large expanses of
▶ Continents land – continents – are an expression of the
greater buoyancy of continental plates compared
to oceanic plates. Continental plates consist of an
upper layer, the ▶ continental crust, which on
Earth is typically 35 km thick, and a lower
Continents layer, called the subcontinental lithospheric man-
tle, which on Earth can reach as deep as 250 km.
Balz Samuel Kamber The density of both layers is lower than that of
Trinity College, Dublin, Ireland typical convecting upper mantle, resulting in a
negative gravity anomaly over continents
(Barrell 1914).
Keywords ▶ Radiogenic isotopes demonstrate that the
two layers are formed contemporaneously during
Buoyancy; Gravity anomaly; Mechanical subduction of oceanic plates (e.g., Nägler
strength; Mountain belt; Plate motion; Radioac- et al. 1997). At depth, where subducted ▶ oceanic
tive heat; Sedimentary basin; Subcontinental lith- crust experiences ▶ metamorphism, hydrous
ospheric mantle; Subduction fluids are released into the mantle, allowing a
partial melt to form, which preferentially con-
sumes the denser mantle minerals and also incor-
Synonyms porates many chemical elements that are not
favored by mantle minerals. This leaves behind
Continental lithosphere; Continental plate; Con- a less dense residue that is depleted in many trace
tinental tectosphere elements, among others, in the radioactive heat
producers. Such subcontinental lithospheric man-
tle acts as a buoyant, strong, cool protective root
Definition for continental crust (Jordan 1978). The melt
itself rises to the ▶ Moho (the boundary between
On a planet with ▶ plate tectonics, ▶ continents the Earth’s crust and mantle) and differentiates
are the topographic expression of emergent, into the continental crust, which is also buoyant
buoyant, strong, and cool plates, contrasting and chemically very different from the bulk
with thinner and hotter oceanic plates covered planet (Taylor and McLennan 1985). The buildup
by water. A relatively thin crust (20–80 km) and of radioactive heat in the crust leads to chemical
an underlying much thicker subcontinental stratification, typically culminating in an event of
▶ mantle (200–300 km) together form the conti- widespread granite emplacement (cratonization)
nental lithosphere that moves as a coherent unit. after which the new crust achieves full mechani-
Together they have the capacity to withstand cal stability (Sandiford and McLaren 2002).
destructive convective forces for billions of On Earth, continents have an average age of
years. The uppermost crust is enriched in many ca. 2.2 billion years; the oldest continental
538 Continuum
fragments are about 3.8 billion years old. Such References and Further Reading
antiquity is testimony to the thermal and mechan-
ical stability of continents (de Wit et al. 1992). Barrell J (1914) The strength of the Earth’s crust. J Geol
22:425–433
The familiar concept of continents surrounded
de Wit M, Roering C, Hart JR, Armstrong RA, de Ronde
by ocean basins may only be valid on planets CEJ, Green RWE, Tredoux M, Peberdy EP, Hart RA
whose interiors cool via the plate tectonic pro- (1992) Formation of an Archaean continent. Nature
cess. Furthermore, as a result of relative plate 357:553–562
Jordan TH (1978) Composition and development of the
motion, the continental plates (which cannot be
continental tectosphere. Nature 274:544–548
destroyed by subduction processes) can thicken McKenzie D, Bickle MJ (1988) The volume and compo-
and deform along their edges, leading to the for- sition of melt generated by extension of the litho-
mation of mountain belts. Continents on young sphere. J Petrol 29:625–679
Nägler TF, Kramers JD, Kamber BS, Frei R, Prendergast
planets with higher crustal heat production have
MDA (1997) Growth of subcontinental lithospheric
less mechanical strength and lower topography, mantle beneath Zimbabwe started 3.8 Ga: a Re-Os
whereas mid-aged planets, like Earth, have con- study on chromites. Geology 25:983–986
tinents capable of supporting rapid vertical move- Rudnick RL (1995) Making continental crust. Nature
378:571–577
ment. Amalgamation of several continental
Sandiford M, McLaren S (2002) Tectonic feedback and
plates produces ▶ supercontinents, whose the ordering of heat producing elements within the
breakup is typically triggered by thermal erosion continental lithosphere. Earth Planet Sci Lett
of the subcontinental lithospheric mantle by heat 204:133–150
Taylor SR, McLennan SM (1985) The continental crust:
upwellings from the deeper mantle (mantle
its composition and evolution, Geoscience Texts.
plumes; McKenzie and Bickle 1988). Blackwell, Oxford
Large sedimentary basins that accommodate
erosion products of mountain belts either express
the thinning of the underlying continental litho-
sphere (rift basins and passive margins), thermal
doming followed by erosion and subsequent
cooling (intraplate basin), or flexural loading of Continuum
lithospheric margins (e.g., foreland basins).
Daniel Rouan
See Also Meudon, France
▶ Continental Crust
▶ Heat Flow, Planetary Definition
▶ Mantle The continuum is the smooth, continuously vary-
▶ Mantle Plume, Planetary ing portion of an ▶ electromagnetic spectrum,
▶ Metamorphism with no spectral features such as atomic or molec-
▶ Moho ular lines or bands. It may be produced by differ-
▶ Oceanic Crust ent physical processes: radiative recombination
▶ Plate, Lithospheric of electrons previously in free states, two-photon
▶ Plate Tectonics decays of metastable levels, thermal bremsstrah-
▶ Radioactive Heating lung, ▶ blackbody radiation, and synchrotron
▶ Radiogenic Isotopes emission.
Convection, Stellar 539
See Also outer third of the radius. This zone is a remnant

from the Sun’s pre-main-sequence epoch, when it
▶ Background was entirely convective.
▶ Blackbody Convection occurs through upwelling of gas
▶ Bremsstrahlung Radiation cells. The hot cells are buoyant and rise toward
▶ Electromagnetic Spectrum the surface, expanding and releasing energy by
▶ Radiative Processes radiation. The cooled cell, now denser, falls back C
down, where it is reheated by the star’s energy
source. The cycle then repeats itself in a circulat-
ing pattern of motion. Convection only occurs if
the specific entropy (entropy per unit mass) of the
Convection, Stellar gas declines toward the surface. In regions where
the entropy rises, fluid cells can rise temporarily,
Steven W. Stahler but then quickly fall back down and thereafter
Department of Astronomy, University of oscillate. Net energy transport in such regions
California, Berkeley, CA, USA occurs through radiative diffusion.
The specific entropy inside a star falls if the
object is either heated from below or cooled from
Definition above. The first mechanism operates in main-
sequence stars, where hydrogen fusion generates
Convection is the transport of heat by turbulent energy in the central region. The second mecha-
motion of gas. Within stars, this thermal energy nism operates in pre-main-sequence stars. These
is supplied either by nuclear fusion or bulk grav- stars have such large radii that surface cooling
itational contraction. The luminosity of the Sun alone drives convection throughout the interior.
is carried outward by convection in the outer The upwelling of convective eddies in the Sun
third of its radius. The shifting pattern of granu- is visible directly in the ever-changing pattern of
lation visible on the solar surface represents granulation near the surface. It is a combination
rising and falling gas cells in the convection of this upward motion and the Sun’s internal
zone. The upwelling of gas in this zone, together rotation that creates the star’s magnetic field.
with the Sun’s rotation, amplifies and maintains This field, in turn, channels the solar wind and
the solar magnetic field. When the Sun was a provides a braking mechanism for the star’s rota-
▶ pre-main-sequence star, its relatively high tion. This general idea is confirmed by observa-
surface cooling drove convection throughout tions of other main-sequence stars. Those with
its interior. outer convection zones rotate much more slowly
than stars lacking them.
The rise and fall of cells, while it has an overall
Overview order, is also a somewhat chaotic process. In this
regard, convection is similar to other forms of
Stars generate energy by nuclear fusion or grav- fluid turbulence, all of which are difficult to
itational contraction. This energy is transported treat in a quantitative manner. The most success-
through the interior, eventually streaming out- ful approach is the mixing-length theory. Models
ward as radiation from the surface. The transport of stellar interiors that use the mixing-length the-
occurs either through the diffusion of radiation or ory accurately reproduce the chief properties of
convection. In the Sun, whose luminosity stems main-sequence stars, including the depth of the
entirely from fusion, convection operates in the convection zone itself.
540 Cool Early Earth
See Also Hadean Earth and that Earth cooled rapidly

from its “hot phase” during and soon after core
▶ Main Sequence, Star formation.
▶ Pre-main-Sequence Star
▶ Protostars
▶ Stellar Winds Overview
The chemical composition of the 4.4–4.3 Ga Jack

Hills zircons implies that they crystallized from
References and Further Reading granitic magma. To form the granite in turn
necessitates that water was transported into the
Foukal PV (2004) Solar astrophysics. Wiley-VCH,
mantle, perhaps by subduction of hydrated oce-
Weinheim
Shore SN (2003) The tapestry of modern astrophysics. anic crust. This implies that oceans were present
Wiley, New York at the surface and that ▶ plate tectonics operated.
The presence of oceans, in turn, allows specula-
tion that life was present at that time. Metamor-
phosed sedimentary rocks and ▶ pillow lavas in
3.8 Ga-old strata from the ▶ Isua Supracrustal
Cool Early Earth Belt (West Greenland) provide evidence of liquid
water and temperate conditions in the early
Nicholas Arndt Archean.
ISTerre, Université Grenoble Alpes, France It is commonly accepted that the ▶ Sun was
less luminous in the first part of Earth history,
25–30 % less than today during the Hadean and
Synonyms early Archean. This should have resulted in
extremely cold conditions at the surface of the
Early Earth Earth and other terrestrial planets. The probable
mechanism to generate temperate surface condi-
tions is the atmospheric presence of a high pro-
Definition portion of greenhouse gases such as CO2 and
methane, combined with some sort of internal
The term “Cool Early Earth” refers to the new regulation that maintained their concentrations
vision of the ▶ Hadean-early Archean Earth at the levels needed to preserve clement condi-
characterized by much more temperate surface tions at the surface (the so-called Faint Young
conditions than believed before. The term was Sun Paradox).
coined by the American geochemist John Valley
in 2002. A popular image of the early Earth is
indeed of a planet covered by hot magma. This See Also
image is at the origin of the term Hadean (from
the Greek god of the underworld, Hades) for the ▶ Archean Eon
first eon. The fractionated oxygen isotopic com- ▶ Earth’s Atmosphere, Origin and Evolution of
positions of 4.4–4.3 Ga-old Jack Hills zircons ▶ Earth, Formation and Early Evolution
indicate, however, that they crystallized from ▶ Faint Young Sun Paradox
magma that had been in contact with water or ▶ Hadean
sediments; thus, surface temperatures may have ▶ Isua Supracrustal Belt
been low enough to permit liquid water on the ▶ Jack Hills (Yilgarn Craton, Western Australia)
Earth’s surface. It is therefore possible that liq- ▶ Oceans, Origin of
uid water was present at the surface of the ▶ Water, Delivery to Earth
Coordinate Systems 541
Synonyms
Coonterunah Subgroup, Australia
Reference frame
Nicholas Arndt
Definition
C
Definition An astronomical coordinate system is a set of
three orthogonal directions, an origin, and a
The Coonterunah Subgroup is one of the oldest choice of mathematical coordinates within this
well-preserved supracrustal sequences known on system, with respect to which the position and
Earth. Located in the ▶ Pilbara Craton, Western motion of celestial bodies are referred.
Australia, the sequence is about 5 km in strati-
graphic thickness and exposed over about 75 km
strike length. It consists mainly of tholeiitic and Overview
minor komatiitic basalt, some felsic lava with inter-
calated carbonate, and ▶ chert beds. It is 3.52 Ga Motion and position are not absolute concepts
old and its metamorphic grade is only mid- and must be described with respect to some ref-
greenschist to lower amphibolite facies. It contains erence. In astronomy, this is a reference frame,
sedimentary carbonate and ▶ kerogen that may that is to say the realization of a set of axes with
represent some of the oldest traces of life on Earth. the means to assign coordinates to an object.
Since these axes are not given a priori on the
celestial sphere, one must use existing celestial
See Also bodies or directions to define the coordinate sys-
tem. In this context, it is important to draw atten-
▶ Archean Traces of Life tion to the difference between a reference system
▶ Basalt and a reference frame. A reference system is the
▶ Chert set of prescriptions stating how a celestial coor-
▶ Kerogen dinate system is to be formed. It defines the origin
▶ Komatiite and fundamental planes (or axes) of the coordi-
▶ Mafic and Felsic nate system, together with the constants and
▶ Pilbara Craton models necessary to fully define the system.
A reference frame consists of a set of identifiable
points on the sky, together with their coordinates,
which serves as the practical realization of a
Coordinate Systems reference system.
A system of axis is determined by the choice
François Mignard of an origin and of three mutually orthogonal
CNRS, Observatoire de la Côte d’Azur, directions labeled by the unit vectors e1, e2, and
University of Nice Sophia-Antipolis, Nice, e3. With the underlying assumptions of absolute
France Euclidean space, the orientation of the system
does not change when translated between two
origins, meaning that the choice of the origin
Keywords and directions are independent of each other.
The commonest origins are the location of the
Astrometry; Declination; Fundamental astron- observer (topocentric frame), the center of the
omy; Hipparcos; ICRF; ICRS; Right ascension; Earth (geocentric frame), the barycenter of the
VLBI solar system (barycentric frame), and the center
542 Coordinate Systems
of the Galaxy (the galactic frame). The center of a the Sun, in its yearly path around the celestial
planet or a satellite can be used as well for spe- sphere, crosses the equatorial plane moving
cific purposes. The set of three directions is from south to north. The ▶ declination (d) of
constructed by selecting the polar direction e3, a body is its angular distance north or south of
or equivalently by selecting a fundamental plane the equator, and the ▶ right ascension (a) is its
going through the origin, and a direction e1 in this angular distance measured eastward along the
plane for the x-axis. A point M is then represented equator from the equinox. Because both the
by the Cartesian coordinates in this frame or by celestial equator and the ▶ ecliptic are moving
spherical coordinates. The latter are more conve- (precession of the equinoxes), the coordinate
nient to represent directions on the celestial systems that they define must be specified at a
sphere and are widely used in observational particular date. In contrast, the ICRF
astronomy or to construct stellar catalogues and (International Celestial Reference Frame) has
ephemerides of low accuracy. Cartesian coordi- fixed directions determined by the positions of
nates are preferred to model motions in the solar a set of extragalactic sources observed in radio
system and to produce accurate ephemerides with interferometry, with an origin at the
position and velocity vectors. The choice of the barycenter of the solar system. However,
fundamental directions is dictated by the neces- these axes correspond closely to what would
sity to make them accessible to observation, so conventionally be described as the equator and
that the system is materialized in space. This equinox of J2000.0 and therefore to the equa-
leads to the following usual systems, appearing torial system at this epoch. For application
in most astronomical work. with accuracy requirement less than 0.1 arcsec,
the distinctions between ICRS and the con-
• The horizon coordinate system uses the ventional definition are not significant. The
observer’s local vertical as a fundamental subset of the single stars of the Hipparcos
direction or equivalently the local horizon as Catalogue is a secondary realization at optical
a fundamental plane. This is a physical defini- wavelengths and is designated as the HCRF
tion that can be accessed with a plumb line or a (Hipparcos Celestial Reference Frame).
spirit level. The pole overhead is the zenith, • Of less importance are the ecliptic system,
and the one diametrically opposite is the nadir. using the mean orbital plane of the Sun as
The two angles that specify the spherical coor- fundamental plane with origin at the vernal
dinates are the azimuth angle (A) and the alti- equinox, and the galactic coordinate system
tude (h). The azimuth is measured in the defined by arbitrarily assigning equatorial
horizontal plane from one of the two points coordinates for the pole and the origin in the
of intersection of the celestial meridian with galactic equator, nominally close to the galac-
the horizon. The altitude is the angular height tic center. In the ICRF, the galactic pole is near
of a point with respect to the horizon, counted ap = 192.75 , dp = 27.13 , and the galactic
positively above the horizon and negatively center is located at about ap = 266.40 ,
below. Instead of altitude h, its complement dp = 28.93 .
measured from the zenith, the zenith distance
z, is also frequently used.
• In the equatorial coordinate system, the pri- See Also
mary direction is defined by the Earth’s spin
axis, giving the ▶ celestial equator (the pro- ▶ Astrometry
jection on the sky of the Earth’s equator) as ▶ Celestial Equator
fundamental plane. The reference point on the ▶ Declination
celestial equator is the vernal equinox, the ▶ Ecliptic
direction on the celestial sphere at which ▶ Right Ascension
Core, Planetary 543
References and Further Reading ▶ Planet Formation

Green RM (1985) Spherical astronomy. Cambridge Uni- ▶ Runaway Gas Accretion
Kovalevsky J, Seidelman PK (2004) Fundamentals of
astrometry. Cambridge University Press, Cambridge
Léna P, Rouan D, Lebrun F, Mignard F, Pelat D (2012)
Observational astrophysics, 3rd edn. Astronomy and C
Astrophysics Library, Springer, Berlin, Chap. 4
Lissauer JJ, Stevenson DJ (2007) Formation of giant
planets. In: Reipurth B, Jewitt D, Keil K (eds) Pro-
tostars and planets V. University of Arizona Press,
Tucson, pp 591–606, 951 pp
Pollack JB, Hubickyj O, Bodenheimer P, Lissauer JJ,
Core Accretion, Model for Giant Podolak M, Greenzweig Y (1996) Formation of the
Planet Formation giant planets by concurrent accretion of solids and gas.
Icarus 124:62–85
Sean N. Raymond
CNRS, Universite de Bordeaux, France
Core, Planetary
Definition
Tilman Spohn
Core accretion is the standard model for giant Deutsches Zentrum f€ur Luft- und Raumfahrt
planet formation. The core accretion model pro- (DLR), Institut f€ur Planetenforschung, Berlin,
poses that the formation of ▶ giant planets starts Germany
with core formation followed by gas accretion.
First, small bodies continually collide to form
larger ones, eventually reaching the stage of pro- Keywords
toplanetary cores, which are essentially large
▶ planetary embryos that form in the giant planet Planets; Satellites
region. Cores that reach masses of a few Earth
masses begin to accrete gas from the ▶ proto-
planetary disk, slowly at first, and then at a run- Synonyms
away rate when the gaseous envelope’s mass
becomes comparable to the core mass. Cores, Planetary core
therefore, represent the “seeds” of gas giant and
ice giant planets in this model. An alternative
model for giant planet formation is disk instabil- Definition
ity that proposes that giant planets form via a
local gravitational collapse in the disk, which The core is the central spherical region of a
can later be followed by core formation and ▶ planet. In a ▶ terrestrial planet, it consists of
accretion of solids. an iron-rich alloy. In a satellite, the core may be
composed of a mixture of iron, silicates, and ice
depending on the degree to which the satellite
See Also interior is differentiated (see ▶ Differentiation,
Planetary). In a ▶ giant planet, the core is made
▶ Atmosphere, Primitive Envelope of a similar mixture of iron, ▶ rock, and ice albeit
▶ Giant Planets at much larger pressures and temperatures.
544 Core, Planetary
Overview The gravity fields measured at the Galilean satel-

lites of ▶ Jupiter suggest that ▶ Io, ▶ Europa, and
The interiors of planets and satellites are differen- ▶ Ganymede have iron-rich cores (e.g., Schubert
tiated to varying degrees with the heaviest mate- et al. 2004). This is particularly true for Ganymede
rials at the center. The degree of differentiation in for which the magnetic field suggests a liquid iron-
solid planets and satellites depends on their ther- rich core with possibly a solid inner core. ▶ Cal-
mal histories since differentiation requires at least listo and ▶ Titan (Iess et al. 2010), as the data
partial melting at some early epoch. Cosmochem- suggest, are incompletely differentiated and likely
ical and physical models of the planets and have large cores composed of ice, silicates, and
satellites suggest the existence of cores and pro- possibly iron or iron oxides. The gravity fields of
vide estimates of their sizes and masses. An iron- Jupiter and ▶ Saturn suggest cores consisting of
rich core has been proven beyond doubt for the ice, silicates, and iron, but their masses and, in
▶ Earth by the inversion of seismic and gravity particular, their radii are uncertain (Guillot and
data. The Earth’s core is composed of a Fe-Ni Gautier 2015 for a review). Estimates range from
alloy but also contains a small fraction (<10 %) a few to about ten Earth masses. Uranus and
of lighter elements such as Si, O, or S and is ▶ Neptune may have similar cores, but interpreta-
layered with a liquid outer core and a solid inner tions of the gravity field suggest that the layering
core. (See Rabinowicz et al. (2015) for reviews on in the interiors of the latter two planets is less
the interior structure of the Earth and Olsen and pronounced than in Jupiter and Saturn. Rather,
Schubert (2015) for reviews on the Earth’s core density and composition change with depth more
dynamics.) It is widely agreed that the inner core is gradually (Podolak et al. 1995).
the result of core freezing and that it grows as the
planet cools. It is also widely agreed that the
energy liberated upon core freezing powers the See Also
generation of its ▶ magnetic field through a
dynamo. ▶ Mars’ core is inferred from gravity ▶ Callisto
data and the chemistry of the ▶ SNC meteorites ▶ Differentiation, Planetary
(Sohl and Schubert 2015 for a review). Its density, ▶ Dynamo, Planetary
chemistry, and the absence of a present-day mag- ▶ Earth
netic field suggest that the core is completely ▶ Europa
liquid (Schubert and Spohn 1990). The cores of ▶ Ganymede
▶ Venus and ▶ Mercury are constrained by the ▶ Giant Planets
average densities of these planets and cosmo- ▶ Io
chemistry. Venus is thought to have a core approx- ▶ Jupiter
imately as large as the Earth’s. The large average ▶ Magnetic Field
density of Mercury along with the recent gravity ▶ Mars
and libration data gathered by the Messenger mis- ▶ Mercury
sion suggests that the core is unusually large with a ▶ Neptune
radius of about 0.8 planetary radii or about ▶ Planet
1,800 km (Smith et al. 2012) and a solid inner ▶ Rock
core. Recent reanalysis of Apollo seismic data by ▶ Satellite or Moon
Weber et al. (2011) suggests that the Moon has a ▶ Saturn
solid inner core with a radius of 240 10 km ▶ Silicate Minerals
surrounded by a fluid, iron-rich outer core of ▶ SNC Meteorites
330 20 km. The interpretation of the data sug- ▶ Terrestrial Planet
gests a partially molten zone in the rock mantle ▶ Titan
above the core of 480 15 km. A solid mantle ▶ Uranus
and a crust complete the lunar interior structure. ▶ Venus
Corona, Coronae 545
References and Further Reading surrounding a conductor, for example, a mineral

surface, which occurs when the strength of the
Guillot T, Gautier D (2015) Giant planets. In: Spohn T, electric field exceeds a certain threshold value.
Schubert G (eds) Treatise on geophysics, vol 10.
A charged ▶ plasma is created around the surface
Elsevier, Amsterdam, pp 529–557
Iess L, Rappaport NJ, Jacobson RA, Racioppa P, Steven- generating ions, which dissipate their charge to
son DJ, Tortora P, Armstrong JW, Asmar S (2010) proximal areas of lower potential or recombine to
Gravity field, shape, and moment of inertia of Titan. form neutral molecules. If the ionized region C
Science 12:1367–1369
continues to grow instead of quenching, a
Olsen P, Schubert G (eds) (2015) Core dynamics, vol 8,
Treatise on geophysics. Elsevier, Amsterdam, 339 pp momentary spark or a continuous arc may result.
Podolak M, Weizman A, Marley M (1995) Comparative A neutral atom or molecule of the medium, in a
models of uranus and neptune. Planet Space Sci region of strong electric field, may be ionized, for
43:1517–1522
example, by the absorption of a photon, to create
Rabinowicz B, Dziewonski A, Schubert G (eds)
(2015) Seismology and the structure of the earth, a positive ion and a free electron. The electric
vol 1, Treatise on geophysics. Elsevier, Amsterdam, field around the surface may then separate these
852 pp charged particles and prevent their recombina-
Schubert G, Spohn T (1990) Thermal history of Mars and
tion. As a result of the acceleration of the elec-
the sulphur content of its core. J Geophys Res
95:14095–14104 trons, further electron/positive-ion pairs may be
Schubert G, Armstrong JD, Spohn T, McKinnon WB created when they collide with other neutral
(2004) Interior composition, structure, and dynamics atoms. These then undergo the same processes
of the Galilean Satellites. In: Bagenal F, Dowling TE,
creating a cascade of electrons.
McKinnon WB (eds) Jupiter. The planet, satellites, and
magnetosphere. Cambridge University Press, Cam- Such energetic processes may have contrib-
bridge, pp 281–306 uted to prebiotic synthesis, providing electrical
Smith D et al (2012) Gravity field and internal structure of energy similar to the electric discharge used in
Mercury from MESSENGER. Science 336:214.
Miller-Urey-type experiments. Corona dis-
doi:10.1126/science.1218809
Sohl F, Schubert G (2015) Interior structure, composition charges are estimated by some to discharge
and mineralogy of the terrestrial planets. In: Spohn T, some three times the energy of lightning strikes
Schubert G (eds) Treatise on geophysics, vol 10. on the present Earth, and this ratio may have been
similar on the primitive Earth.
Weber RC et al (2011) Seismic detection of the lunar core.
Science 331:309. doi:10.1126/science.1109375
See Also
Corona Discharge ▶ Miller, Stanley

▶ Plasma
Institute of Technology, Meguro–ku, Tokyo, Japan
Institute for Advanced Study, Princeton, NJ, USA Corona, Coronae
Washington, DC, USA Jörn Helbert
Center for Chemical Evolution, Georgia Institute DLR, Institut f€ur Planetenforschung, Berlin,
of Technology, Atlanta, GA, USA Germany
A corona discharge is a positive or negative elec- Coronae are volcano-tectonic landforms only
trical discharge created by the ionization of a gas seen on the planet ▶ Venus. They are oval to
546 Coronagraphy
circular features typically 100–300 km in diame-

ter. A few of the coronae are even larger. They
have a circular or nearly circular, tectonically
deformed annulus, which usually stands a few
hundred meters above the surrounding plains.
The area inside the annulus is typically lower
than the surrounding plains and has been flooded
with plains-forming volcanic lava. Aprons of
young lobate (tongue-shaped) volcanic flows are
seen radiating from many coronae. At the center
of some coronae one finds the corona core, an
elevated and tectonically deformed area.
See Also
▶ Venus
Coronagraphy
Daniel Rouan Coronagraphy, Fig. 1 Bernard Lyot, a French astrono-

LESIA, Observatoire Paris-Site de Meudon, mer who invented the coronagraph, an optical instrument
Meudon, France whose original purpose was to study the faint extended
emission around the Sun, the so-called corona
Definition the glare of a star to allow detection of compan-

ions or disk structures in its immediate vicinity.
Coronagraphy designates the group of optical
techniques that aim at suppressing or reducing
the halo of light that surrounds the image of a Overview
star, in order to detect faint structures like a
circumstellar disk or companions, especially The ability to directly image an extrasolar
exoplanets. It was invented in 1930 by the French planet – that is, to separate the light emitted or
astronomer Bernard Lyot (Fig. 1) to study the reflected by its surface or atmosphere from that of
Sun’s corona, the extremely faint emission from the star it orbits – offers the greatest prospect for
the region around the Sun, at times other than characterizing these objects. Direct ▶ imaging
during a solar eclipse. The coronagraph is in its gives the possibility of determining the colors
simplest form an occulting disk in the focal plane and spectra of planets, in a way independent of
of a telescope combined with a mask in a plane their orbital inclination. This information allows
optically conjugated to the entrance aperture, so astronomers to distinguish between gas giants,
that the image of the solar disk is blocked and the ice giants, and Earth-like planets under a variety
stray light reduced by a large factor. Since then, of circumstances such as distance from the star,
the term has been kept for research in areas other age, etc. Direct imaging also allows astronomers
than the solar corona and designates now any to study the planet’s atmosphere, possible cloud
optical system able to block as much as possible systems, etc., and may also become the
Coronagraphy 547
successful way for establishing the habitability of (Dl/l = 10 %) or the spectroscopic study of
an exoplanet. possible exoplanet atmospheric features at
Directly imaging extrasolar planets requires various different wavelengths.
(a) angular resolution sufficient to spatially sepa- • Imperfections in the telescope mirrors and coat-
rate the planet from its central star and (b) the ings cause speckles in the image, that is, small
means to suppress the diffracted light from the spots of extra light intensity, after the
central star such that the planet’s brightness coronagraphic mask; their intensity and vari- C
becomes comparable to or greater than the resid- ability can easily be high enough to obscure the
ual diffracted star light. extremely faint exoplanet image. It is possible
Concerning the first point, at optical wave- to reduce these speckles using deformable mir-
lengths a conventional telescope of several rors (▶ Adaptive Optics), controlled thanks to
meters diameter is in principle sufficient to spa- smart algorithms, or to code them in order to
tially resolve a planet on an orbit comparable to recognize them from the planet image.
those observed in our solar system, around nearby • Finally, the weak signal from an extrasolar
stars. However, the telescope must be in space or planet requires substantial integration times
equipped with an extremely efficient ▶ adaptive (many hours) during which it is essential that
optics system; otherwise, the blurring due to the residual stellar leakage be kept extremely
atmospheric turbulences at ground level and at stable, which is especially challenging on the
high altitude would degrade too severely the ground, considering atmospheric as well as
image quality. thermal fluctuations. This stresses the need to
As regards the second requirement, this is conduct research of Earth-sized exoplanets
precisely the role of a coronagraph – to block from space, but, even then, drifts and vibra-
the starlight as efficiently as possible, using opti- tions in the telescope cause varying stellar
cal elements within the telescope. “speckles” that can limit the ultimate sensitiv-
The main challenge of using an optical coro- ity for planet detection.
nagraph for exoplanet imaging is the star-planet
brightness contrast ratio. For an Earth-sized Under the pressure of the planet hunters,
planet orbiting a star similar to our Sun, it is coronagraphy is an area that has shown enormous
about 1010–1 in the visible. Any successful star- progress in the past decade as presented below,
light suppression technique must reduce this con- with a flourishing of very new concepts and tech-
trast ratio by a huge factor and must do so at a nical breakthroughs. Ground-based instruments
very small angle, typically equal to y = 2–5 l/D, are already capable of direct detection of
where l is the wavelength of observation and D is exoplanets in the most favorable cases, and a first
the diameter of the telescope (this value corre- generation of coronagraphic instruments installed
sponds to the fundamental limit imposed by the on the largest telescopes (8–10 m diameter) will
phenomenon of ▶ diffraction). An important soon extend by a large factor the population of
parameter is thus the smallest star-planet angular directly detected exoplanets. It is rather likely that
separation at which this suppression level is within another decade or so, a coronagraphic space
achieved to avoid the need for primary mirror mission will be able to obtain the first images of
diameter larger than, say, 10 m. planets that are analogs to our Earth.
There are other challenges albeit a little less
stringent.
Basic Methodology
• The optical bandwidth of the light suppression
technique – that is, the range of wavelengths A star observed with a perfect telescope in space,
over which the required light suppression can thus unaffected by the aberrations of the Earth’s
be maintained – should be as large as possible atmosphere, will produce an image still
to enable either broadband photometry surrounded by a halo of light ▶ diffracted from
548 Coronagraphy
into a vast family of devices that can be broken

down into various broad categories: amplitude or
phase-mask coronagraphs, interferometric corona-
graphs, and pupil apodization. A fifth family is the
external occulter. Each one offers pros and cons
that can be measured by a few key parameters.
Four of those parameters are especially important:
• The inner working angle (IWA) is the angle of

separation from the star below which the flux
from the planet is rapidly attenuated and/or
beyond which the flux from a resolved star
Coronagraphy, Fig. 2 The airy pattern is the appear- rapidly increases. Currently, the most mature
ance that the image of a star made by a perfect telescope in concepts feature an IWA of 4 l/D, and it is
space would present. Because of the phenomenon of dif- suggested that an IWA of 2 l/D is
fraction, it is not an extremely sharp spot but rather a
achievable.
central core surrounded by characteristic rings. The larger
the diameter of the telescope and the smaller the wave- • The planet throughput, that is, the fraction of
length, the sharper the diameter of the core the planet’s light that reaches the detector.
Current throughputs of 8–30 % are measured,
the edges of the telescope aperture. Known as the but claims that a throughput of 80 % or higher
Airy pattern, this halo, which is structured with is reachable are made. In addition, several
concentric rings of decreasing intensity (see coronagraphs produce stellar halo suppression
Fig. 2), is many orders of magnitude brighter within only a fraction of the entire field of
than any extrasolar planet image that would be view, for instance, a sector or a ring, some-
superimposed on it. Suppressing this halo is the times referred to as the “discovery space.”
role of a coronagraph. • The sensitivity of the coronagraph to wavefront
errors such as image position and focus. These
The Basic Lyot’s Coronagraph defects arise due to drift and misalignment;
The main finding of Bernard Lyot (1939) is that even very small ones and their effects can
blocking the light with an opaque mask in the hide or mimic a planetary signature.
focal plane that follows the contour of the bright • The chromaticity of the coronagraph, that is,
object (the Sun’s surface in his case) is not suffi- its ability to suppress starlight across a broad
cient at all, because light diffracted by the edge of wavelength range, thus allowing a lower total
the mask will spread far beyond the limits of the integration time. Typical useful bandwidth is
beam defined by geometric optics and finally will Dl/l 20 % or less.
be able to refocus. This is illustrated in Fig. 3 (top
and mid panels): the first mask in the focal plane A thorough study on how each coronagraph
blocks the core of the starlight but leaves residual design behaves with respect to those key param-
diffracted light in the reimaged pupil (blue eters was done by Guyon et al. (2006).
beam). Lyot realized that if a mask with a central
circular hole slightly smaller that the pupil image Improvement on the Lyot Concept with
was installed, then it would block the essential Amplitude Masks
part of this diffracted light as shown on the bot- Those concepts evolved from Lyot’s original
tom panel of Fig. 3. solar coronagraph, using a mask at an image of
the star, and another mask at a later image of the
The New Coronagraphic Concepts entrance aperture. It can be shown that, in this
Over the past decade, coronagraphic concepts basic form, Lyot’s concept cannot reach the
have proliferated far beyond Lyot’s basic vision rejection required for planet detection by several
Coronagraphy 549
Coronagraphy, Fig. 3 The principle of the Lyot’s coro- the image. Middle: An opaque mask is put at the focal
nagraph. Top: optical scheme of an imaging system with plane and blocks a large part of the light; however,
relay optics. The image of a point-like object at infinity because of diffraction by the edges of the mask, a fraction
(star) is formed by a telescope (a) at its focal plane (b); a of the light is spread beyond the geometrical beam and
lens at this location forms the image of the telescope produces a significant intensity in the focal plane. Bottom:
aperture in another plane (c) where a third lens a ring mask placed in the pupil plane is efficient to block
re-images the star in plane D. The intensity of the light most of this spurious light
distribution is represented by the curve which is a cut of
orders of magnitude. An important variant of this entrance pupil with a dedicated transmission
concept is the band-limited coronagraph mask (Soummer 2004). Laboratory demonstra-
(Kuchner and Traub 2002), where the focal tions to date have achieved contrast levels in the
plane mask features a carefully tailored transmis- 106 range. One advantage is that it may be easily
sion profile confining the residual diffracted light adapted to conventional telescopes with central
to a well-defined region of the pupil plane. This obstruction, such as those used in the next gener-
type of coronagraph has produced excellent lab- ation of ground-based coronagraph projects.
oratory results with a measured contrast of
6 1010, albeit only in monochromatic light. Phase-Based Lyot Coronagraphs
Another Lyot variant is the Apodized-Pupil A similar approach to the Lyot concept is to
Lyot Coronagraph (APLC), which modifies the induce a phase shift rather than an amplitude
550 Coronagraphy
φ=π φ=0
φ=0 φ=π
4QPM
Mask
Stop
Coronagraphy, Fig. 4 Schematics describing the principle of the four-quadrant phase-mask coronagraph and the
distribution of the amplitude of the light at the various planes of the optical system
variation in the focal plane; destructive interfer- rejection performance increasing exponentially
ence is then produced for on-axis starlight at the with the number of devices. A recently proposed
pupil level. In the first proposed idea (Roddier variant of the 4QPM is the optical vortex corona-
and Roddier 1997), the opaque spot of Lyot was graph (Foo et al. 2005), which uses a spiral-
simply replaced by a transparent spot of a proper staircase phase mask. They are more robust
size. The best-known example of this concept against degradation due to the star’s position.
remains the four-quadrant phase-mask (4QPM)
coronagraph (Rouan et al. 2000), which focuses Pupil Apodization
the starlight onto a mask that shifts light passing Using the Fresnel principle, it can be shown that
through half the focal plane by half a wavelength the diffraction pattern at the focus of a
(see Fig. 4). Such coronagraphs have near-ideal telescope – that is, the actual distribution of inten-
theoretical performance with high throughput sity in the image of a point source at infinity, such
and an IWA of 1 l/D. However, they are also as a star – is directly related to the shape of the
sensitive to star position in the focal plane with entrance aperture. More precisely, this intensity
respect to the center (angle y), with performance is given by a mathematical transformation of the
degrading rapidly with tip/tilt errors. One variant aperture, called the Fourier transform. The sharp
(Rouan et al. 2007) that allows improving on this edges of a conventional telescope result in the
sensitivity is the 8-quadrant phase mask, where Fourier ringing that produces the well-known
the canceling function is proportional to y4 rather Airy pattern. Conceptually, the simplest corona-
than y2. There exist several solutions to manufac- graph would entail a modification of the tele-
ture them for broadband light: one uses polariza- scope aperture so it lacks these sharp edges.
tion properties of uniaxial crystals, and another is Called apodization, this modification can be
based on the stacking of several 4QPM with achieved by tapering the telescope transmission
Coronagraphy 551
Coronagraphy, Phase induced amplitude apodization

Fig. 5 The principle of the
phase-induced-amplitude-
apodization coronagraph
that produces apodization
by modifying the light
intensity within the beam
C
PIAA entrance pupil
Focusing element
PIAA exit pupil
Focal plane
through a semitransparent mask that falls gradu- redistribute the uniform beam of light into a
ally to zero transmission at the edges. These tapered profile (see Fig. 5). The off-axis planet
masks trade throughput for inner working angle, image is highly distorted, but this effect can be
with a typical mask designed for an IWA of 4 l/D compensated. Contrary to classical apodization,
having a transmission of 8 %. However, no light collected by the entrance pupil is lost,
manufacturing such a mask with graded attenua- and the inner working angle can be as small as
tion is extremely difficult. Significant effort has 2–3 l/D for a throughput of 80 % or more. How-
been invested in binary approximations to these ever, this requires optical surfaces very difficult to
apodization functions, using sharp-edged metal manufacture, located in various conjugate planes.
masks that suppress diffraction over only part of
the field of view (Kasdin et al. 2003). Referred to Interferometric Coronagraph
as “shaped pupils,” these masks, which can The nulling coronagraph is a coronagraph based
include micron-sized features, must be carefully on a nulling interferometer, as opposed to more
fabricated but are feasibly produced with current familiar designs of an apodized aperture tele-
manufacturing techniques. These designs are scope and Lyot coronagraph. This family uses
inherently achromatic and may be attractive for concepts of ▶ interferometry and aims at produc-
characterization of planets with known positions, ing destructive interference between two beams.
where they can be optimized over a narrow The achromatic interferometric coronagraph
region of the focal plane. Shaped-pupil masks (AIC) was the first of this type to be proposed,
have been demonstrated in the laboratory at the by Gay and Rabbia as early as 1997, and has been
2 109 contrast level, even in broadband light. refined since then (Rabbia et al. 2007). It is basi-
An alternative approach to performing cally a Michelson-Fourier interferometer modi-
apodization is to modify the phase of the light to fied by inserting on one arm an achromatic p
modulate the intensity in the pupil plane. The phase shift and a pupil rotation by 180 (see
phase-induced-amplitude-apodization corona- Fig. 6). The smart idea is to produce the p phase
graph (Guyon 2003) of O. Guyon is based on this shift by crossing of a focus, thus making it per-
approach, using highly aspheric mirrors to fectly achromatic. The collimated beam from the
552 Coronagraphy
Coronagraphy,
Fig. 6 The principle of the
achromatic interferometric Coronagraphic output
coronagraph (AIC) based
on destructive interference
between two beams, one
having suffered the
crossing of a focus Cat’s eye system Beam recombiner
Beam splitter
None coronagraphic output

3 flat mirrors
Collimated light beam

Piezotranslator used for that feeds the AIC
the zero OPD setting
telescope is split into two sub-beams forming the recently done led to some mission scenarios that
two interferometric arms, one where the focus- yield satisfactory efficiency with one occulter and
crossing occurs. The beams are recombined after- much better with two occulters. The deployment of
ward. Other designs have been proposed, such as the large starshade and the propulsion required for
the nulling coronagraph of M. Shao (2007). many stellar observations remain major technol-
ogy issues to be addressed.
External Occulter Coronagraphs
This type of coronagraph belongs to a peculiar
family, since it relies on optical systems separated Future Directions
by several 10, 000 km! In 1960, Lyman Spitzer
proposed the combination of a telescope and a Wavefront Control
starshade in space for discovery of planets. The A problem common to all internal coronagraphs is
concept implied enormous distances and starshade wavefront accuracy and stability. Wavefront
dimensions. With recent studies (Cash 2006), the errors produced by imperfections in the telescope
inter-spacecraft separation has been reduced and mirrors and coatings cause speckles in the image
the level of suppression improved by using petal after the coronagraph masks have suppressed most
shapes that would permit operation using a shade of the starlight. The intensity and variability of
with a nominal diameter of 40 m at a telescope- these speckles can easily make the faint exoplanet
starshade separation of 40,000 km (see Fig. 7). The image indiscernible. Achieving Airy ring suppres-
telescope can be an ordinary space telescope, and sion sufficiently to allow detection of Earth-sized
its diameter is determined mainly by the require- planets would require wavefront phase errors of
ment to detect faint planets. It has inherently ach- less than an angstrom (1010 m) – well beyond
romatic properties, a strong advantage. However, current polishing capabilities for telescope-sized
operation remains the main drawback, since mirrors. However, the use of a smaller deformable
pointing from one star to another requires that the mirror in the optical train allows irregularities in
starshade travel several thousand kilometers. To the wavefront to be corrected by feedback, up to a
accomplish this within a few weeks demands large spatial frequency set by the deformable mirror’s
starshade velocities and accelerations and thus a actuator spacing (Bordé and Traub 2007). With
substantial power. However, the engineering effort state-of-the-art deformable mirrors, this allows a
Coronagraphy 553
Coronagraphy, Fig. 7 An artist view of the “New conventional space telescope. The insert shows a simu-
Worlds Imager” project based on a large deployable exter- lated image of the solar system as it would be detected
nal occulter at a distance of 40,000 km from a
large fraction of low- and mid-spatial-frequency such as a Fourier transform of the image. This is,
errors to be removed from the wavefront, produc- for instance, the principle of the self-coherent
ing an image with a characteristic “dark hole” camera proposed by Baudoz et al. (Galicher and
central region (see Fig. 8). It can be shown that Baudoz 2007).
with two deformable mirrors, both phase and
amplitude defects can be corrected, for an even Differential Coronagraphy
better improvement. Even with the best correction of the wavefront,
there will still remain some defects, and it
Calibration of Residual Speckles becomes clear that a good way of correcting
The field image is small, a few arcsec in diameter, them is to subtract a similar image that presents
but an average flux of 1010 times the star flux the same defects, for instance, of another star or
implies that there could be several hundred the same star, but where presumably no planet
speckles as bright as an Earth-like planet. Only image would be present. Several concepts have
one of those speckles is a planet. One technique to been proposed in that spirit and several of them
identify it makes use of the coherence of starlight, have been tested on the sky: spectral differential
that is, light from the star is coherent (i.e., will imaging, rotational differential imaging, and
interfere coherently) with speckles whose origin polarization differential imaging.
is scattered starlight, but will not interfere with In spectral differential imaging (Racine
light in the focal plane that comes from the planet et al. 1999), two images of the star are taken at
(or dust) orbiting the star. Because of the inter- two close wavelengths, carefully chosen so that
ferences, the intensity distribution of the stellar the planet would have a different contrast, for
flux features fringes that striate the speckles. The instance, within and outside a spectral feature
fringes can be detected by a proper processing, characteristic of some expected compound in its
554 Coronagraphy
Coronagraphy, Fig. 8 A deformable mirror with many actuators (left) can provide a fair correction of the residual
defects in the wavefront, producing an image with a characteristic “dark hole” central region (right)
atmosphere (e.g., methane). In another version λmax

(Sparks and Ford 2002), one produces the spectra
of all the pixels in the image, with a so-called
integral field spectrograph, and looks for differ-
ences in behavior between speckles, whose size
changes with wavelength, and a planetary image
whose size is constant, as illustrated in Fig. 9.
Rotational differential imaging (Marois
et al. 2006) requires that the telescope be rotated
a few degrees around its axis several times, an
image being taken at each position: the optical
defects will rotate as well, in sky coordinates,
while the planet image would remain stable. Fig-
ure 10 illustrates this technique. λmin
Finally, in polarization differential imaging,
one makes the reasonable guess that the light
θ=0 Planet
from the planet is significantly polarized, while
the starlight is not: images taken in different
Coronagraphy, Fig. 9 Integral field spectroscopy pro-
polarization states could reveal the planet after duces spectra of all points in an image: most of the light
subtraction. that remains after coronagraphy is due to speckles whose
size changes with wavelength (l is along the vertical axis),
while a planet would be of about a constant size with
Combining All Techniques
wavelength
The challenge to beat a contrast of 1010 is so
difficult that it becomes more and more evident
that the good strategy cannot be to rely on one will pick up the first image of a cousin of our
technique alone, but to combine many of them. Earth, will have included together an efficient
The coming years, or decades, will probably coronagraph combining pupil and image plane
show that the winning instrument, the one that masks, a super-polished telescope with no central
CoRoT 7b 555
Coronagraphy, Fig. 10 Rotational differential imaging. The telescope is rotated around the line of sight and images
are taken at each angle. The optical defects that rotate with the telescope can be subtracted
obstruction, one or two deformable mirrors, sev- Lyot B (1939) The study of the solar corona and promi-
eral differential imaging techniques, and most nences without eclipses (George Darwin lecture,
1939). Mon Not R Astron Soc 99:580
probably will have been operated in space. Marois C, Lafrenière D, Dovon R, Macintosh B, Nadeau
D (2006) Angular differential imaging: a powerful
high-contrast imaging technique. Astrophys
See Also J 641:556–564
Rabbia Y, Gay J, Rivet J-P (2007) The achromatic
▶ Adaptive Optics interfero coronagraph. CR Phys 8:385
Racine R, Walker GAH, Nadeau D, Doyon R, Marois
▶ Diffraction
C (1999) Speckle noise and the detection of faint
▶ Direct-Imaging, Planets companions. Publ Astron Soc Pac 111:587–594
▶ Exoplanets, Discovery Roddier F, Roddier C (1997) Stellar coronagraph with
▶ Imaging phase mask. Publ Astron Soc Pac 109:815–820
Rouan D, Riaud P, Boccaletti A, Clénet Y, Labeyrie
A (2000) The four-quadrant phase-mask coronagraph
References and Further Reading I. Principle. Publ Astron Soc Pac 112:1479–1486
Rouan D, Baudrand J, Boccaletti A, Baudoz P, Mawet D,
Bordé P, Traub W (2007) Speckle noise reduction tech- Riaud P (2007) The four quadrant phase mask corona-
niques for high-dynamic range imaging. CR Phys graph and its avatars. CR Phys 8
8:349 Shao M (2007) Calibration of residual speckle in a nulling
Cash W (2006) Detection of Earth-like planets around coronagraph. CR Phys 8:340
nearby stars using a petal-shaped occulter. Nature Soummer R (2004) Apodized pupil Lyot coronagraphs for
442:51 arbitrary telescope apertures. Astrophys
Foo G, Palacios DM, Swartzlander GA (2005) Optical J 618:161–164
vortex coronagraph. Opt Lett 30:3308–3310 Sparks WB, Ford HC (2002) Astrophys J 578:543
Galicher R, Baudoz P (2007) Expected performance of a
self-coherent camera. CR Phys 8:333
Guyon O (2003) Phase-induced amplitude apodization of
telescope pupils for extrasolar terrestrial planet imag-
ing. Astron Astrophys 404:379 CoRoT 7b
Guyon O, Pluzhnik EA, Kuchner MJ, Collins B, Ridgway
ST (2006) Theoretical limits on extrasolar terrestrial Nader Haghighipour
planet detection with coronagraphs. Astrophys J Suppl
Ser 167:81
Kasdin NJ, Vanderbei RJ, Spergel DN, Littman MG Manoa, Honolulu, Hawaii, HI, USA
(2003) Extrasolar planet finding via optimal apodized
pupil and shaped-pupil coronagraphs. Astrophys
J 582:1147–1161
The exoplanet CoRoT-7b is the seventh planet
Kuchner M, Traub W (2002) A coronagraph with a band-
limited mask for finding terrestrial planets. Astrophys detected via transit photometry (Fig. 1) by the
J 570:900–908 European space telescope, CoRoT (Léger
556 CoRoT 7b
The 153 transits of CoRoT − 7b et al. 2009). With a radius of ~1.6 Earth radii, this
1.0015 planet is the first potentially terrestrial-like planet
1.0010 found outside our solar system. CoRoT-7b
revolves around its central star in a very short-
Realtive Flux
1.0005
period orbit of 20 h making this planet one of the
1.0000 few with extremely small orbital periods. The
0.9995
close proximity of this planet to its Sun-like
host star (which is about 500 light-years away)
0.9990 implies that it must experience extreme condi-
0.9985 tions. For instance, its surface may be covered
−0.2 −0.1 0.0 0.1 0.2 by lava or boiling oceans (Fig. 2).
Time (days) Since the discovery of CoRoT-7b, many
attempts have been made to determine the mass
CoRoT 7b, Fig. 1 Light curve of CoRoT-7 showing
153 transits of CoRoT-7b of this planet using radial velocity data (Léger
et al. 2009; Queloz et al. 2009; Hatzes et al. 2010,
2011; Boisse et al. 2011; Ferraz-Mello
et al. 2011). However, due to stellar activities,
the value of the mass of this planet is uncertain.
The current estimates put the mass of this planet
in the range of 2–8 Earth masses (Fig. 3).
Discovery of CoRoT-7b opened a new field in
the studies of extrasolar planets. The mass of this
planet, although uncertain, puts this object in the
category of super-Earths. The latter motived
many researchers to develop models for the inte-
rior and composition of super-Earth planets.
Results suggested that despite the shaky value
of its mass, CoRoT-7b has probably a rocky inte-
rior and internal composition similar to that of
Earth.
Being so close to its host star, CoRoT-7b may
CoRoT 7b, Fig. 2 Artist’s rendering of CoRoT-7b not have significant atmosphere. Because this
covered with lava planet is in a tidally locked position with its
CoRoT 7b, Fig. 3 CoRoT-

7b in comparison to the
solar system planets
CoRoT Satellite 557
parent star, which causes one side of it to face the Overview

star at all time. It is believed that the surface of its
bright side must have been depleted of volatiles, CoRoT (for Convection, Rotation, and Planetary
suggesting that the atmosphere of the planet may Transits) is the first space mission designed for
be rich with rocky vapors. the study of the stellar physics through
▶ asteroseismology and for the search for
exoplanets with the method of transits. It was C
launched on December 27, 2006, from
References Baı̈konour, and sent scientific data up to Novem-
ber 2012. CoRoT was developed and operated by
Boisse I et al (2011) Disentangling stellar activity and the French space agency ▶ CNES and French
planetary signals. Proc Int Astron Union 273:281
CNRS laboratories LESIA, LAM, IAS, and
Ferraz-Mello S et al (2011) On planetary mass determina-
tion in the case of super-Earths orbiting active stars. IRAP, with significant contributions from Aus-
The case of the CoRoT-7 system. Astron Astrophys tria, Belgium, Brazil, ESA, Germany, and Spain.
531:A161
Hatzes AP et al (2010) An investigation into the radial
velocity variations of CoRoT-7. Astron Astrophys
520:A93 Basic Methodology
Hatzes AP et al (2011) On the mass of CoRoT-7b.
Astrophys J 743:75 Stars, acted on by gravity, pressure, and Coriolis
Léger A et al (2009) Transiting exoplanets from the
forces, behave as oscillators with many specific
CoRoT space mission VIII. CoRoT-7b: the first
super-Earth with measured radius. Astron Astrophys modes. These oscillations are detectable through
506:287 tiny variations in the stellar luminosity, and their
Pont F, Aigrain S, Zucker S (2010) Re-assessing the analysis provides important parameters on the
radial-velocity evidence for planets around CoRoT-7.
stars’ internal structure.
Mon Not R Astron Soc 411:1953
Queloz D et al (2009) The CoRoT-7 planetary system: two When an exoplanet travels between its host
orbiting super-Earths. Astron Astrophys 506:303 star and us, it induces a weak drop of the star’s
luminosity (a mini-eclipse). The detection of this
periodic ▶ transit in the star’s light curve,
together with its careful analysis, reveals the
existence of the exoplanet and provides important
CoRoT Satellite characteristics such as its radius, its orbit param-
eters, and the rotation period of the star.
Olivier La Marle CoRoT’s wide field of view enables it to mon-
Centre National d’Etudes Spatiales DSP/EU, itor the flux from thousands of stars for
Paris, France uninterrupted period of several months, in order
to detect and measure the tiny variations in this
flux, either for stellar activity measurements or for
Keywords exoplanets’ transit detection. The satellite carries a
27 cm, two mirrors off-axis visible telescope focus-
Asteroseismology; CoRoT; Exoplanet; Transit ing the stars’ light on a focal unit hosting the four
(1) frame transfer CCD matrices of 2,048 4,096
pixels (Baglin et al. 2007; Auvergne et al. 2009).
Definition The detectors, whose temperature is regulated at
40 C, have a high quantum efficiency in the
The CoRot mission is a satellite for the study of visible wavelengths. On top of this instrument, a
the stellar structure through asteroseismology baffle reduces the parasitic stray light down to a
and for the detection of transiting ▶ exoplanets. few photons/s/ per pixel (Fig. 1).
558 CoRoT Satellite
Sun and larger than expected. In terms of stellar

structure, the first seismic interpretations of the
measured eigen-frequencies address the crucial
question of the extension of the mixing beyond
the stellar convective core. This key process is
responsible for the present large uncertainty on
stellar age determinations.
Red Giants and the Future of Our Sun

Toward the end of their lives, stars like the Sun
expand and become ▶ red giant stars. Because of
the turbulent convection in their outer layers, red
giant stars are expected to exhibit solar-like oscil-
lations, but in a much lower range of frequencies
(10–100 mHz). CoRoT data allowed to measure
clearly for the first time such oscillations in a
large sample of red giants (Fig. 2).
In addition, unambiguous evidence of the
excitation of both radial and nonradial modes
was provided, previously an open question.
CoRoT data also confirmed the existence of
modes with lifetimes of the order of one month
(De Ridder et al. 2009; Carrier et al. 2010).
The first theoretical modeling of this stellar
evolution stage suggests that it would be possible
CoRoT Satellite, Fig. 1 The CoRoT satellite during its
integration onto the Soyuz launcher to explain the observed oscillation spectra and
their variety for different stages of the structure
of stars along their expansion in the red giant
1. From March 2009, only two CCD matrices phase. Indeed these red giants of different masses
were left operating. and ages are representative of all the successive
generations of stars in the Galaxy. Through the
astreroseismology of red giants, CoRoT therefore
Key Research Findings opened the door for a new method for galactic
archeology.
Asteroseismology
One hundred fifty-six various bright stars and New Type of Pulsators
35,000 red giants were observed for CoRoT data led to the discovery of new types of
uninterrupted durations of about 80–150 days in pulsating stars. HD180872 is one of them. This
most cases. A few highlights of the results star was known to belong to the Beta Cephei class
already achieved by CoRoT are presented here. of pulsators, young massive stars which are pro-
genitors of SN-II type supernovae and thus
Solar-Like Pulsation, Granulation, and Convective mainly responsible for the enrichment of the uni-
Core verse in carbon and oxygen. These stars classi-
CoRoT measured solar-like oscillations and cally show oscillation periods of the order of a
granulation in stars hotter than the Sun (Michel few hours. In the lightcurve of HD180872,
et al. 2008). The modes’ widths (inversely pro- CoRoT data revealed, at very low amplitude,
portional to the lifetime of the modes) have been the existence of higher-frequency modes, due to
found to be noticeably larger than those in the stochastic oscillation, very comparable to the
CoRoT Satellite 559
CoRoT Satellite, 15,000

Fig. 2 Power spectrum of
the CoRoT observations of
the red giant HR 7349
Power [ppm2]
(Carrier et al. (2010). 10,000
Reproduced with
permission # ESO)
C
5,000
0
0 10 20 30 40 50
Frequency [μHz]
CoRoT Satellite, 1.0002

Fig. 3 CoRoT-7b light
curve. The 0.03 % dip in the 1.0001
star’s flux during the
planet’s transit is clearly 1.0000
visible (# CoRoT)
DF/F
0.9999
0.9998
0.9997
0.9996
−4 −2 0 2 4
Time (h)
ones observed in the Sun (Belkacem et al. 2009). frontier with ▶ brown dwarfs (Deleuil
This confirms the existence of a powerful con- et al. 2008). On the opposite side, CoRoT-7b is
vective zone and will allow the scaling of its the first telluric exoplanet which mass and radius
energy. This discovery opens new perspectives are known (Léger et al. 2009; Queloz et al. 2009).
in the study of these objects where low-frequency It provides evidence for the existence of Earth
oscillations and high-frequency ones could be cousins in orbit around Sun-like stars. Several
used in a complementary way to probe the center planets such as CoRoT-1b, CoRoT-2b, CoRoT-
and the outer layers of the star. 11b, and CoRoT-20b challenge the current theory
for planet formation and/or migration in a disk.
Search for Exoplanets CoRoT-7b is phase-locked, that is to say that
One hundred sixty three thousand six hundred its orbital and rotational periods are equal
eighty two stars were monitored by CoRoT for (0.85 day). Therefore, it shows always the same
duration up to 6 months. Thirty-five companions face to its star (as the Moon does relatively to the
(mainly exoplanets and a few brown dwarves) Earth). This induces a very hot temperature (up to
have been confirmed so far, and 130 candidates 2,600 K) on the dayside and a very cold one
identified (Fig. 3). (down to 50 K) on the nightside.
These discoveries widen the variety of the CoRoT-9b is the first transiting temperate
known exoplanets family (Table 1). CoRoT-3b, exoplanet. Indeed, the medium surface tempera-
the heaviest and the densest one, lies at the ture of this Jupiter-like gaseous planet stands
560 CoRoT Satellite
CoRoT Satellite, Table 1 Characteristics of a sample of CoRoT planets

Mean
Period Mass Radius density Star
Name (day) (MJupiter) (RJupiter) (g/cm3) type Main features
CoRoT-1b 1.51 1.03 1.49 0.38 G0V Metal poor host start – secondary transit
detected
CoRoT-2b 1.74 3.31 1.46 1.31 G7V Very young star (~0.5 billion years),
challenges planet formation
theory – secondary transit detected
CoRoT-3b 4.26 21.6 1.0 26.4 F3V Brown dwarf or super planet
CoRoT-5b 4.03 0.46 1.39 0.217 F9V Very low density
CoRoT-7
b 0.85 0.025 0.157 7.50 G9V CoRoT-7b first telluric exoplanet
c 3.7 0.043 NA NA (M = 5.4 MEarth, R = 1.6 REarth)
d 9 NA NA NA
CoRoT-9b 95.27 0.84 1.05 0.90 G3V First moderate temperature transiting
exoplanet
CoRoT-10b 13.24 2.75 0.97 3.7 K1V High eccentricity (0.53)
CoRoT-11b 2.99 2.33 1.43 0.99 F6V Fast-rotating star (period<2 days)
CoRoT-14b 1.51 7.6 1.09 7.3 F9V Very dense hot giant
CoRoT-15b 3.06 63.3 1.12 96 F7V Very dense brown dwarf
CoRoT-20b 9.24 4.24 0.84 8.9 G2V Very young system (100 million years).
A very dense planet challenging planet
formation theories. High eccentricity
(0.56)
between 20 and 150 C, depending on the (detected for the first time for a substellar com-
models, and the temperature excursion between panion with CoRoT-3b).
daytime and nighttime is probably low (Deeg
et al. 2010). Its low eccentricity ensures a rela-
tively low seasonal temperature variation. Fur- Applications
ther spectroscopic observations of its transits
should open a new era in the study of exoplanets, CoRoT will let us make a giant step forward in
allowing planetologists and biochemists to enter our knowledge of stars’ and the planets’ birth,
the game. life, and death and therefore bring a major con-
CoRoT also detected the secondary transit of tribution of our knowledge of the Universe we
exoplanets at the visible wavelength for the first live in. It also paves the way for the search for life
time (Alonso et al. 2009a, b). This phenomenon in the universe: on one hand, it identifies planets
occurs when the planet disappears behind its star, for which detailed analysis with other techniques
inducing a slight dip in the total flux received will be of particularly high interest. On the other
from the two of them. In the case of CoRoT-1b, hand, it eased the work of its successors, the US
for instance, this dip was only of 2/10,000. This mission ▶ Kepler (Basri et al. 2005) and the
allowed to measure the planet albedo future European mission ▶ Plato (Catala
(reflectivity), which turns to be of about 10 % et al. 2010), by providing a great know-how on
(compared to about 40 % for the Earth). the transit method, in particular on the optimal
In addition to transits, CoRoT detected a num- strategy for targets selection, for data processing,
ber of other effects and demonstrated how they and on-the-ground follow-up. It also helps the
could be used to better characterize the star- delicate thinking of the astronomical community
planet systems, such as the Rossiter-McLaughlin in the definition of the best strategy for the search
effect and the ellipsoidal and beaming effects for extraterrestrial life.
CoRoT Satellite 561
Future Directions ▶ Hot Jupiters

▶ Hot Neptunes
In stellar physics, CoRoT was a pioneer in ▶ James Webb Space Telescope
establishing the richness of asteroseismology. ▶ Kepler Mission
The future will probably be devoted to the ▶ Microlensing Planets
increase of the sample and of the variety of stars ▶ Planetary Migration
observed. CoRoT, Kepler, and later on Plato will ▶ PLATO 2.0 Satellite C
be major contributors; ground surveys might also ▶ Radial Velocity
contribute, especially for low-frequency ▶ Red Giant
oscillations. ▶ Secondary Eclipse
Several international scientific bodies were set ▶ Spectroscopy
up to propose a roadmap for the exoplanets and ▶ Stellar Pulsation
extraterrestrial life detection, in Europe ▶ Stellar Rotation
(EP-RAT, Blue Dots initiative) or in the USA ▶ Super-Earths
(Decadal Survey). The Pathways Conference in ▶ Transit
Barcelona in September 2009 proposed a three- ▶ Transiting Planets
step strategy: ▶ VLT
1. Statistical study of planetary objects

2. Designate sources suitable for spectroscopic References and Further Reading
follow-up
Alonso R, Guillot T, Mazeh T et al (2009a) The secondary
3. Carry out spectroscopic characterization
eclipse of the transiting exoplanet CoRoT-2b. Astron
The first and second steps would involve sev- Alonso R, Alapini A, Aigrain S et al (2009b) The second-
eral methods such as transits, radial velocities, ary eclipse of CoRoT-1b. Astron Astrophys
506:331–336
astrometry, and microlensing. The Kepler mis-
Auvergne M, Bodin P, Boisnard L et al (2009) The CoRoT
sion and, on a larger scale, the Plato mission satellite on flight: description and performances.
intend to make a wide census and characteriza- Astron Astrophys 506:411–424
tion of transiting planetary systems. TESS, Baglin A et al (2007) The CoRoT mission and its scientific
objectives. AIP Conf Proc 895:201–209
CHEOPS, GAIA, Euclid, and ground surveys
Basri G, Borucki WJ, Koch D (2005) The Kepler mission:
will also contribute. The spectroscopic charac- a wide-field transit search for terrestrial planets. New
terization will require large ground telescopes, Astron Rev 49(7–9, Spec Iss):478–485
such as the ESO VLT and the future large Belkacem K, Samadi R, Goupil MJ et al (2009) Solar-like
oscillations in a massive star. Science
telescopes (E-ELT, TMT), and space-based
324(5934):1540–1542
facilities such as the James Webb Space Bouchy F, Queloz D, Deleuil M et al (2008) Transiting
Telescope. exoplanets from the CoRoT space mission III. The
spectroscopic transit of CoRoT-Exo-2b with SOPHIE
and HARPS. Astron Astrophys 482:25–28
Carrier F, De Ridder J, Baudin F et al (2010) Non-radial
See Also oscillations in the red giant HR 7349 measured by
CoRoT. Astron Astrophys 509:A73
▶ Asteroseismology Catala C, Arentoft T, Fridlund M et al (2010) PLATO:
planetary transits and oscillations of stars – the
▶ Atmosphere, Temperature Inversion
exoplanetary system explorer. ASP Conf Ser
▶ Brown Dwarf 430:260–265
▶ Exoplanets, Discovery De Ridder J, Barban C, Baudin F et al (2009) Non-radial
▶ Exoplanets, Modeling Giant Planets’ oscillation modes with long lifetimes in giant stars.
Nature 459:398–400
Atmospheres
Deeg HJ, Moutou C, Erikson A et al (2010) A transiting
▶ Habitability, Effect of Eccentricity giant planet with a temperature between 250 K and
▶ Habitable Zone, Effect of Tidal Locking 430 K. Nature 464:384–387
562 Corotation Torque
Deleuil M, Deeg H, Alonso R et al (2008) Transiting

exoplanets from the CoRoT space mission-VI. Corrosion
CoRoT-Exo-3b: the first secure inhabitant of the
Brown-dwarf desert. Astron Astrophys
491(3):889–897 ▶ Oxidation
Deleuil M, Bonomo AS, Ferraz-Mello S et al (2012)
Transiting exoplanets from the CoRoT space mission.
XX. CoRoT-20b: a very high density, high eccentricity
transiting giant planet. Astron Astrophys 538:A145
Léger A, Rouan D, Schneider J et al (2009) Transiting Cosmic Background Radiation
exoplanets from the CoRoT space mission-VIII.
CoRoT-7b: the first Super-Earth with measured radius. Stéphane Le Gars
Mazeh T, Faigler S (2010) Detection of the ellipsoidal and Centre François Viète, Université de Nantes,
the relativistic beaming effects in the CoRoT-3 Nantes, France
lightcurve. Astron Astrophys 521:L59
Michel E, Baglin A, Auvergne M, Catala C et al (2008)
CoRoT measures solar-like oscillations and granula-
tion in stars hotter than the Sun. Science Keywords
322(5901):558–560
Queloz D, Bouchy F, Moutou C et al (2009) The CoRoT-7 Arno Penzias; Astronomy; Big Bang; Cosmic
planetary system: two orbiting super-Earths. Astron background radiation; Cosmology; George
Gamow; Georges Lemaı̂tre; History; Nucleosyn-
thesis; Physics; Robert Wilson
Corotation Torque Synonyms
Sean N. Raymond CMB; Cosmic microwave background

History
Definition In 1965, two American physicists, Arno Penzias

and Robert Wilson, accidentally discovered elec-
The corotation torque is that exerted on a planet’s tromagnetic background noise at 7.35 cm wave-
orbit by material (usually gas) co-orbiting with length in the millimeter radio wave region.
the planet. This has a very important effect on the Penzias and Wilson were working for the Bell
orbital migration of planets of less than roughly Telephone Laboratories on satellite communica-
50 Earth masses that form in gaseous ▶ proto- tion improvement when they brought to light this
planetary disks. Recent analysis suggests that homogeneous and isotropic radiation resulting
very rapid, inward “type 1” migration – caused from the universe as its whole; it was quickly
by tidal interaction between the disk and a read as fossil radiation from a universe that was
planet – may in fact be slowed or even reversed originally dense and hot and had cooled off
in certain situations due to the corotation torque. because of its expansion to a 3.5 K temperature
today. In 1978, Penzias and Wilson got the Nobel
Prize for this discovery.
See Also However, there had been some previous inves-
tigations in this general area. The French physi-
▶ Lindblad Resonance cist Charles-Edouard Guillaume had calculated
▶ Planetary Migration in 1896, by Abney’s works and according to the
▶ Protoplanetary Disk black body’s law that Stefan granted in 1879, that
Cosmic Ray in the Galaxy 563
an isolated body in the space, only subdued to Marc L-R (2004) Cosmologie scientifique. Rev Métaphys
stars’ radiation, would see its temperature De Morale 43:399–411
Penzias AA, Wilson RW (1965) A measurement of excess
increasing of nearly 5 K. In 1926, Sir Arthur antenna temperature at 4080 Mc/s. Astrophys
Eddington shows that the real temperature of J 142:419–421
the space, due to star’s radiance, is 3.18 K. But Wilson Robert (1978) The cosmic background radiation.
there was a lack of theoretical framework to give Nobel Lecture, 8 décembre
sense to theoretical calculations assuming some C
model of the universe. Indeed, Edwin Hubble
proved in 1924, with detailed observations, that Cosmic Dust
the universe is formed of myriads of galaxies. In
1929, he also discovered empirically that these ▶ Interstellar Dust
galaxies all move away from one another. At the
same time, the works of the Belgian physicist
Georges Lemaı̂tre interpreted the observational Cosmic Microwave Background
data in terms of relativity theory to show that the
universe is expanding. ▶ Cosmic Background Radiation
In the 1940s, the American astrophysicist
George Gamow laid down the basis of nucleosyn-
thesis, the formation of the chemical elements. Cosmic Ray in the Galaxy
According to Gamow, the chemical elements
could have been made at a hot and dense period Nikos Prantzos1 and Jun-Ichi Takahashi2
1
of the universe, in the bosom of an “original soup” Institut d’Astrophysique de Paris, Paris, France
2
that he named “ylem” and that represented an NTT Microsystem Integration Laboratories,
early stage of the Big Bang. In 1948, two Atsugi, Japan
Gamow’s collaborators, Alpher and Herman,
developed Gamow’s ideas and predicted the exis-
tence of a diffuse radiation background with a 5 K Keywords
temperature. Within this theoretical framework,
this cosmic background radiation has become the Cosmic rays; Electrons; Energetic particles;
experimental mainstay of the Big Bang theory, Nuclei
which includes the universal expansion and the
nucleosynthesis of the lightest elements. The
background radiation has recently been measured Definition
very accurately in 2001 and 2006, respectively,
with satellites COBE and WMAP. Galactic cosmic rays are high-energy
(relativistic) electrons and nuclei, accelerated by
supernova explosions and massive stellar winds
References and Further Reading and traveling through the Galaxy by scattering on
fluctuations of interstellar magnetic fields, which
Acker A (2005) Astronomie-Astrophysique-Introduction. render their flux isotropic.
Dunod, Paris
Assis AKY, Neves MCD (1995) The redshift revisited.
Astrophys Space Sci 227:13–24
Charles-Edouard G (1896) La température de l’espace. La Overview
Nat 24:234
Dicke RH, Peebles PJE, Roll PG, Wilkinson DT The energy spectrum of galactic cosmic rays
(1965) Cosmic black-body radiation. Astrophys
J 142:414–419
(CR) covers the energy range from a few
Eddington AS (1926) The internal constitution of the MeV/nucleon up to 1015 MeV/nucleon and is
universe. Cambridge University Press, Cambridge well approximated by a power-law N(E) / Ea
564 Cosmic Ray, Ionization Rate
of slope a = 2.7 below 106 GeV/nucleon and ▶ Evolution, Biological

a = 3 above that. Below a few GeV/nucleon, ▶ HZE Particle
the CR spectrum progressively flattens (with a ▶ Ionizing Radiation, Biological Effects
even becoming positive) and its intensity varies, ▶ Mutation
in a way that anticorrelates with solar activity ▶ Nucleon
(“solar modulation”); the solar wind prevents ▶ Proton Irradiation
the lowest energy CR from entering the helio- ▶ Radiation Biology
sphere. The total galactic power of CR is esti- ▶ Radiochemistry
mated to be 1041 erg/s, that is, about 10 % of the ▶ Spallation Reaction
total kinetic power of ▶ supernovae and stellar ▶ Supernova
winds, which are thought to be the main CR
accelerators. However, no galactic accelerator
can account for the highest energy CR (up to References and Further Reading
1012 GeV), the origin of which remains
Strong A, Moskalenko I, Ptuskin V (2007) Cosmic ray
unknown. The residence time of CR in the galac- propagation and interactions in the Galaxy. Annu Rev
tic disk is 107 years. The composition of CR Nucl Part Sci 57:285–327
nuclei is overall similar to the solar one: it con-
sists of 87 % protons, 12 % alpha nuclei
(helium atoms), and 1 % heavier nuclei
(in addition to the nucleonic component 3 % Cosmic Ray, Ionization Rate
of the CR flux are high-energy electrons). The
major uncertainties are at extremely high ener- Steven B. Charnley
gies and for elements heavier than iron. There are Solar System Exploration Division, Code 691,
several important differences: the metallicity (see Astrochemistry Laboratory, NASA Goddard
▶ Metallicity) of CR is 10 times solar, with Space Flight Center, Greenbelt, MD, USA
refractory elements (Fe-peak nuclei) relatively
more enhanced than the volatiles; and the fragile
Li, Be, and B nuclei are 106 times more abundant Definition
in CR than in the Sun. These features suggest that
CR are accelerated from a mixture of interstellar The cosmic ray ionization rate in the ▶ interstel-
gas and dust grains (where refractory elements lar medium is the rate at which H2 molecules as
are overabundant), and during their propagation well as atomic H and He are ionized by the flux of
in the Galaxy they spallate abundant CNO nuclei galactic cosmic rays. This process is fundamental
to produce Li, Be, and B. to driving the chemistry in dense interstellar
From the standpoint of astrobiology, CR may clouds. Current estimates place the value in the
play a crucial role, either as effective energy range 1–40 1017 s1 with the higher values
sources for synthesizing the precursors of terres- being more appropriate for lower-density diffuse
trial bioorganic compounds in interstellar media or interstellar clouds.
as agents inducing mutations in living organisms
and thereby promoting biological evolution. They
may even be lethal in case of a nearby supernova. See Also
▶ Cosmic Ray in the Galaxy

See Also ▶ Cosmic Rays in the Heliosphere
▶ Diffuse Cloud
▶ Biostack ▶ Interstellar Chemical Processes
▶ Cosmic Rays in the Heliosphere ▶ Molecular Cloud
Cosmic Rays in the Heliosphere 565
Overview
Cosmic Rays in the Heliosphere
Cosmic radiation observed at the Earth consists
Don F. Smart of 83 % protons, 12 % alpha particles, 1 %
Air Force Research Laboratory (Emeritus), heavy nuclei with atomic number >2, and 3 %
Bedford, MA, USA electrons. The cosmic ray composition of the
most common elements has been measured to a C
reasonable precision (see Fig. 1). The major
Keywords uncertainties are at extremely high energies and
for elements heavier than iron. The local inter-
Cosmic radiation; Solar modulation stellar spectrum (outside the heliosphere) is con-
stant, but inside the heliosphere, the spectrum and
fluence of particles below 10 GeV/nucleon are
modified by solar activity with a phase that is the
Definition inverse of the solar sunspot cycle. The cosmic ray
flux decreases with increasing energy and the
Cosmic rays are very energetic particles thought proton flux is reduced by 50 % at 1.5 GeV. The
to pervade space. Within the solar system, the cosmic ray flux of the heavier nuclei is reduced
cosmic ray flux is modulated by the solar activity by 50 % at 0.9 GeV/nucleon. The maximum total
cycle. isotropic flux in free space at 1 AU is 3 particles
Cosmic ray spectrum

Phi = 500
100 H
He
C
10−1 H O
Mg
He
10−2 Ne
N
B
Flux(m2 s sr MeV/n)−1
10−3 C,O Li
Be
N Al
10−4 Si
Fe Na
10−5 Fe
Ca
Cr
10−6 Ti
Fe
Mn
10−7 Ar
Ni
10−8
K
Cl
P
10−9 V
S
Sc
10−10 Co
101 2 3 102 2 3 103 2 3 104 2 3
Energy (MeV/n)
Cosmic Rays in the Heliosphere, Fig. 1 A model of during average low solar activity conditions. The elements
the modulated differential cosmic ray spectrum at 1 AU listed on the right side of the figure are in the order of their
for the most abundant elements in the cosmic ray flux observed abundance. Original figure created by the author
566 Cosmic Spherules
Cosmic Rays in the Cosmic ray proton spectrum

Heliosphere, Fig. 2 A modulation Phi = 400 to 1,600
model of the modulated 101
differential cosmic ray
proton spectrum at 1 AU for
100
Flux (M2 S Sr MeV/nuc)−1

modulation conditions,
0
ranging from near solar 40
minimum to solar 500
10−1
maximum. Original figure
created by the author 1,000
10−2
1,500 00
1,6
10−3
10−4
101 102 103 104
Energy (MeV/nuc)
cm2 s1 during solar minimum conditions; dur- References and Further Reading
ing solar maximum conditions, the total cosmic
ray flux is reduced by 40 %. The diffusion of the Townsend LW, Badhwar GD, Blakely EA, Braby LA,
Cucinotta FA, Curtis SB, Fry RJM, Land CE, Smart
cosmic ray flux inwards through the turbulent
DF (2006) Information needed to make radiation pro-
interplanetary medium results in an average tection recommendations for space missions beyond
radial cosmic ray gradient of a few percent per low-earth orbit. NCRP Report No. 153, National
AU. The cosmic radiation flux at Mars (1.5 AU) Council on Radiation Protection and Measurements,
Bethesda
is only a few percent larger than at Earth.
The major factors modulating the cosmic radi-
ation intensity include the solar wind speed, tur-
bulence in the solar wind, and solar magnetic
Cosmic Spherules
polarity. Cosmic ray modulation theory ade-
quately models the flux at the Earth and at our
Frank Sohl
most distant space probes. The modulation param-
eter, designated by the symbol w, is normally
expressed in units called MV. The modulation
Germany
parameter at 1 AU during solar minimum typically
ranges between 400 and 500 MV; however,
extreme solar activity can generate transient mod-
Definition
ulation levels in excess of 1,600 MV (see Fig. 2).
The particle flux below 2 GeV/nucleon
Spherule comes from the Greek word “sphaira”
makes the most important contribution to radia-
for sphere. Cosmic spherules are solid, rounded
tion dose. Since the energy deposition in matter is
particles ranging from micrometer to millimeter
proportional to the square of the atomic charge,
in size with distinct chemical compositions indi-
the heavy elements are important for computing
cating their extraterrestrial origin. Cosmic spher-
radiation dose.
ules are predominantly found in ocean floor
deposits on Earth and are mainly produced by
See Also frictional heating, melting, and ▶ ablation of
▶ meteoroids upon atmospheric entry. Solidified
▶ Cosmic Ray in the Galaxy impact-induced molten droplets of meteoroid and
▶ Cosmic Ray, Ionization Rate target materials are less abundant on Earth but
Cosmochemistry 567
would predominate on the exposed surfaces of were revered as gods in many places of the Med-
atmosphere-less bodies like the Moon that were iterranean basin. During the Middle Ages, they
strongly gardened by impacting meteoroids in were still considered with superstition and fear.
their early history. Though more rational interest was paid to mete-
orites in the Renaissance, they still belonged to
the reign of natural wonders. Interestingly, the
See Also Enlightenment era failed to identify the true prov- C
enance of meteorites, probably because analyti-
▶ Ablation cal chemistry was yet to be developed.
▶ Meteoroid Ernst Florens Chladni (1756–1827) can be
▶ Moon, The seen as the founder of cosmochemistry. At a
time when most scientists believed meteorites
were mere stones from volcanoes or atmospheric
condensations, he proposed in a provocative
Cosmochemistry pamphlet published in 1794 that meteorites
were extraterrestrial objects. This bold proposi-
Matthieu Gounelle tion was harshly discussed for the next 10 years
Laboratoire de Minéralogie et Cosmochimie du among the European scientific community. It was
Muséum (LMCM) MNHN USM 0205 - CNRS finally accepted in 1803 after the fall of a mete-
UMR 7202, Muséum National d’Histoire orite at L’Aigle in France and the publication of a
Naturelle, Paris, France detailed report by Jean-Baptiste Biot
(1774–1862, Fig. 1). Being the first scientist to
travel to the place of a meteorite fall and helped
Keywords by a beautiful literary style, Biot was able to
convince his peers that stones fell from the sky.
Accretion disk; CAIs; Chondrites; Chondrules; At that time, many of the early cosmochemists
Protoplanetary disk such as Chladni, Laplace (1749–1827), or Biot
thought meteorites came from the Moon. It is not
until the mid-nineteenth century that scientists
Synonyms realized that most meteorites come from ▶ aster-
oids, the first of which, Ceres, was discovered in
Meteoritics 1801. After the recognition of the extraterrestrial
nature of meteorites, the field of cosmochemistry
blossomed. It started with the development of
Definition elaborate classification schemes (strikingly similar
to the one used now) and chemical analyses, which
Cosmochemistry is the study of the formation and expanded on the pioneering studies of Lavoisier
evolution of the Solar System and its individual (1743–1794) and Howard (1774–1816). The iden-
components through the analysis of extraterres- tification of new minerals and of chondrules, a
trial samples in the laboratory. major component of the primitive meteorites
unknown in terrestrial rocks, clearly set meteorites
apart from terrestrial rocks.
History
The first mention of a meteorite in western liter- Basic Methodology

ature concerns the prediction of the fall of a
meteorite by Anaxagoras of Clazomenae in the Meteorites are studied using the same techniques
year 467 BC. During Antiquity, ▶ meteorites as terrestrial rocks. The petrography (relative
568 Cosmochemistry
Today, extraterrestrial samples can be collected

directly from space thanks to sophisticated and
expensive space missions, such as Stardust,
which brought back cometary dust from the
Wild 2 comet in January 2006. But these missions
are still rare. Most cosmochemists focus their
work on meteorites and micrometeorites that
fall on Earth, a cheaper way to sample celestial
bodies millions of kilometers away. Roughly 50 t
of meteorites (size range 1 g–100 kg) and 7,000 t
of micrometeorites (< mg) fall on the Earth’s
surface every year. In April 2010, the number of
meteorites registered by the Meteoritical Society
was 38,800.
Nowadays, cosmochemistry is a very active
scientific discipline, which greatly benefits from
its proximity with other fields, especially astro-
physics, but also geology and material sciences.
Thanks to the diversity of extraterrestrial samples
(meteorites as well as lunar rocks and cometary
dust brought back by space missions) and to the
wealth of techniques used and its fecund interac-
tion with a large number of neighboring fields, it
Cosmochemistry, Fig. 1 Jean-Baptiste Biot
(1774–1862). After the fall of the L’Aigle meteorite in
can address a diversity of fundamental scientific
1803 in France, he wrote a report which definitely questions. The following point of view on cos-
established the extraterrestrial origin of meteorites mochemistry is complementary to that presented
in the entry ▶ meteorites; special emphasis will
be given to important discoveries and open
abundance and textural relationships between the questions.
different components and individual minerals)
and mineralogy (structure and chemical compo- What Do Meteorites Look Like?
sition of minerals) are made on rock sections (i.e., Meteorites are rocks made of minerals (and occa-
cut and polished) with a diversity of optical and sionally glasses). They can be distinguished from
electronic microscopes. Isotopic analyses are terrestrial rocks mainly by their fusion crust
performed with a diversity of mass spectrome- (▶ Meteorite, Allende), i.e., a mm-thick veneer
ters. Secondary ion mass spectrometers and of glass produced during their entry at cosmic
thermo ionization and inductively coupled velocity (10–20 km/s) into the Earth’s
plasma mass spectrometers are used for a wide atmosphere.
range of rock-forming elements. While the for- Though meteorites can be divided into 135 dif-
mer provide a very good spatial resolution ferent classes, mineralogically there exist three
(micron scale), the latter have reached precisions main groups of meteorites (Fig. 2): stones (94 %
of the order of tens of ppm. Organic matter within of the observed falls), stony irons (1 %), and irons
meteorites is now studied in detail with tech- (5 %). Though some of them might have an
niques imported from the oil industry impact origin, irons are believed to represent the
(▶ Meteorite, Murchison). cores of large, differentiated asteroids or
Cosmochemistry 569
Cosmochemistry,
Howardites
Fig. 2 Simplified
Irons Eucrites
classification of meteorites.
Diogenites
Gray boxes indicate
differentiated meteorites, Ureilites
while white boxes indicate Mesosiderites
primitive meteorites. Stony-irons
Primitive achondrites Angrites
(shaded gray) are objects
Pallasites C
having the composition of Aubrites
chondrites and the texture
of differentiated rocks Lunar
Shergottites
Chassignites
Nakhlites
Differenciated
Winnonaites
Achondrites
Brachinites
Primitive
Acapulcoites
Stones
Lodranites
Chondrites
H
Ordinary L
LL
Rumurutite
EH
Enstatite
EL
Carbonaceous
Cl CM CO CV CK CR CH CB
planetesimals; and stony-iron meteorites are small mm-sized blobs known as chondrules (see
thought to be mixtures of core and mantle or below) that have a chemical composition similar
core and crust material. Stony meteorites can be to that of the Sun. Achondrites are magmatic
divided into chondrites and achondrites. The for- rocks. They lack chondrules and have a strongly
mer are characterized by a large abundance of fractionated chemical composition relative to
570 Cosmochemistry
Cosmochemistry,
Fig. 3 Chemical
composition of the
chondrite GRO 03116. Red,
green, blue, yellow, and
white code, respectively,
for iron, magnesium,
silicium, calcium, and
aluminum. One can
appreciate the richness in
reduced iron (red), the high
abundance of chondrules
(green-blue iron-
magnesium silicate
spheres), and the rarity of
CAIs (in yellow white)
(Picture courtesy of Anton
Kearsley (NHM))
that of the Sun; they are believed to originate fusion crust, they would be difficult to distinguish
from the crust of planets and other smaller differ- from terrestrial basalts, though detailed studies
entiated bodies. show they come from evolved bodies, which
Chondrites are made of calcium-aluminum- had a different geologic history than the Earth.
rich inclusions (CAIs), chondrules, and matrix
(Fig. 3). The relative abundance of these three Where Do Meteorites Come From?
components varies between chondrite groups. In Most meteorites are chondrites. They are cosmic
most chondrites (except CIs), chondrules repre- sediments whose components were made in the
sent the dominant component. CAIs are an protoplanetary disk and cemented together via
assemblage of calcium and aluminum oxides poorly characterized processes. Chondrites were
and silicates (▶ CAIs). Some of them show evi- not differentiated, i.e., metal and silicates were
dence of melting. Chondrules are an assemblage not segregated after melting. The larger a body is,
of iron-magnesium silicates plus metal and sul- the easier it is to melt; therefore, the undiffer-
fides. Chondrules usually show an igneous tex- entiated nature of chondrites indicates they orig-
ture indicating they were at some point inate from small celestial bodies, ▶ asteroids or
extensively melted in the solar protoplanetary ▶ comets. The determination of the orbits of a
disk. Matrix is made of fine-grained (<1 mm) few meteorites has confirmed that most chon-
iron-magnesium silicates, metal, and sulfides. In drites come from asteroids, though some, such
the case of carbonaceous chondrites, the matrix is as CI chondrites, could come from comets
rich in organic matter. For most chondrite groups, (▶ Meteorite, Orgueil). It is however not possible
the matrix has been modified on the parent aster- to pinpoint from which specific asteroid meteor-
oid due to secondary geological processes such as ites come from, except for one noticeable excep-
thermal metamorphism or hydrothermal tion. The good match between the infrared
alteration. spectrum of HED meteorites and the asteroid
Achondrites, at first sight, look like terrestrial (4) Vesta suggests these meteorites come from
igneous rocks. They are made of mafic minerals that asteroid.
such as olivine and pyroxene and an aluminum- On the other hand, the compositional and min-
rich silicate, plagioclase. If it were not for the eralogical similarities of roughly 100 meteorites
Cosmochemistry 571
206
18 Pb, with respective half-lives of 0.7 and
CO2 4.5 Ga. The long-lived radioisotope 87Sr (T1/2
16 N2 = 48 Ga) was also extensively used to date
early Solar System processes during the 1960s
Gas in EET 79001
40
Ar
and 1970s, but is today too imprecise for the
14
task (compared to other systems). Iodine-129
with half-life of 15.7 Ma was the first short- C
12 36
Ar lived radionuclide whose past presence in a mete-
20
Ne orite was unambiguously demonstrated. Many
84
10 Kr more were to follow (see below).
132
Xe The use of chronometers, combined with astro-
8 nomical observations, made it possible to date the
8 10 12 14 16 18 sequence of events from the collapse of the portion
Martian atmosphere of a molecular cloud to the Solar System in its
Cosmochemistry, Fig. 4 Chemical composition of the present configuration. The lifetime of the molecu-
gas trapped in the SNC (Martian) meteorite EET 79001 lar cloud precursor of our Solar System is not
compared to that of the Martian atmosphere determined by known, but molecular clouds usually live for a
the Viking spacecraft in the 1970s (From Bogard few million years before they get disrupted by the
et al. (1984))
harsh effects (photoionization, powerful winds) of
the stars to which they gave birth. The following
to lunar samples collected by the space missions stage, i.e., the gravitational collapse of a portion of
Apollo and Luna establish that they come from the molecular cloud, takes some 100,000 years and
the Moon. The same number of meteorites – can be seen as a transition phase during which,
forming the SNC (Shergottite-Nakhlite- however, important gas-grain chemistry occurs. In
Chassignite) group – come most likely from the the next stage, the protosun is surrounded by an
planet Mars. This was demonstrated by the excel- accretion disk through which it is fed with matter.
lent match in composition between gas bubbles Because it is within the same disk that the compo-
trapped within meteorites from the SNC group nents of chondrites, chondrites themselves, plane-
and the Martian atmosphere composition mea- tesimals (from 1 km up to 1,000 km), and large
sured by the Viking landers in 1976 (Fig. 4). planetary embryos (1,000 km-sized bodies) are
The origin of micrometeorites is still debated. formed, the accretion disk or solar nebula is also
Recent work suggests they come mostly from called a protoplanetary disk.
comets rather than from primitive asteroids. It is CAIs most likely formed first in the protoplan-
worth noticing that the precise origin of micro- etary disk 4,568 Ga ago, followed by chondrules
meteorites does not matter much, as the analysis which formed 1.5 Ma later. The accretion of chon-
of cometary samples returned by the Stardust drites from their components and of planetesimals
mission has demonstrated that comets and prim- from chondrites occurred as soon as chondrules
itive carbonaceous asteroids are similar in nature. were formed. The disk was dissipated after a few
million years, according to astronomical observa-
Dating Meteorites and Other Bodies: A Rough tions of other stellar systems. Embryos probably
Sequence of Events grew on that same million-year timescale.
One of the major achievements of cosmochemis-
try has been to date extraterrestrial samples. In Chemical and Isotopic Composition
1956, Claire Patterson proposed an age of CI chondrites have the same chemical composi-
4.55 Ga for the Solar System and the Earth (see tion as the Sun for all elements except for the
Age of the Earth). To accomplish this fundamen- most volatile ones, such as H, He, C, and N
tal task, he used two long-lived radionuclides (Fig. 5). This discovery suggests that chondrites
235
U and 238U which decay into 207Pb and are extremely primitive rocks that sample the
572 Cosmochemistry
14 exception to the similarity discussed above, in

that it varies a great deal. While chondrules
12
H have a composition roughly similar to that of
He the Earth, CAIs are enriched in 16O by at least
Solar photosphere
10
C O 4 % relative to chondrules. The origin of that
N
8 enrichment – known since 1973 – is not well
6
understood. The most popular model – the self-
shielding model – invokes photochemical pro-
4 cesses either in the parent molecular cloud or in
the protoplanetary disk (▶ Oxygen Isotopes).
2
Li
0 Short-Lived Radionuclides
0 2 4 6 8 10 Short-lived radioisotopes (SRs) are radioactive
Cl chondrites elements with half-lives ranging from a few
Cosmochemistry, Fig. 5 Chemical composition of the weeks to 100 Ma. Their presence in the nascent
Sun’s photosphere compared to that of the CI chondrites Solar System is inferred from excesses of their
(shown on a logarithmic scale, normalized to silicon) daughter isotopes in meteoritic components,
mostly in primitive components such as the
earliest phases of the protoplanetary disk. The CAIs. Some SRs were present in the protoplane-
relatively small chemical fractionation of other tary disk at abundances significantly higher than
chondrites relative to CI chondrites is attributed the expected average contribution of the interstel-
to physical processing in the protoplanetary disk, lar medium. These SRs therefore require a last-
either before or during chondrule formation. minute origin. They were either made within a
The bulk isotopic and chemical composition star such as a supernova and injected into the
of chondrites is the result of the chemical evolu- nascent protoplanetary disk or produced within
tion of the Galaxy over the last 10 billion years. the disk itself via nuclear reactions between solar
Interestingly enough, the isotopic composition of cosmic rays and ambient dust. The origin of SRs
all but a few elements in Solar System bodies is a hotly debated topic of cosmochemistry, as it
(chondrites, achondrites, Earth, Mars, etc.) is has important consequences for our understand-
remarkably identical in meteoritic, lunar, and ing of: (1) the early Solar System chronology,
terrestrial samples. This indicates that Solar Sys- (2) planetesimal heating, and (3) the astrophysi-
tem matter was well homogenized by high- cal context of our Solar System birth.
temperature processes in the protoplanetary If SRs were homogeneously distributed in the
disk, possibly linked to chondrule formation. Solar System, they can be used to define a chro-
A few solids escaped this processing and pre- nology of Solar System events. Aluminum-26
served their chemical and isotopic properties. (T1/2 = 0.74 Ma), 53Mn (T1/2 = 3.7 Ma), and
These are called presolar grains. They were dis- 182
Hf (T1/2 = 9.0 Ma) are especially useful as
covered in the matrix of chondrites in the late chronometers because their initial content is
1980s at the University of Chicago. They were known in a diversity of objects. Ages based on
isolated thanks to severe chemical treatments and SRs confirm the early formation of CAIs relative
recognized because of their isotopic composition, to chondrules and the rapid evolution of Solar
radically different from that of Solar System mat- System bodies. Dating of some iron meteorites
ter. Presolar grains were made in the atmospheres indicate indeed that differentiation of large plan-
of stars formed hundreds of millions before our etesimals or asteroids occurred contemporane-
Solar System. Thanks to their study, stellar nucle- ously with CAI formation.
osynthesis has become an experimental field. Aluminum-26 and 60Fe (T1/2 = 2.6 Ma) emit
The oxygen isotopic composition of meteor- gamma rays when they decay. They have been
itic components, e.g., CAIs and chondrules, is an proposed as heat sources for planetesimals. The
Cosmochemistry 573
extent of heating and therefore the subsequent probably the most abundant solids in the disk.
geological evolution depends on the initial con- Some authors estimate that there might have
tent of these two SRs in the parent-body been between 1024 and 1025 g of chondrules pro-
considered – and therefore on their initial content duced in the asteroid belt. Understanding how
in the Solar System – as well as on the timing of they were made from precursor solids and disk
formation of the bodies considered. Current gas is therefore a key task of cosmochemistry.
models indicate that 26Al is a more efficient heat It is widely believed that chondrules were C
source than 60Fe. heated up to 2,000 K on timescales of a few
The presence of 60Fe in the Solar System seems minutes and cooled relatively slowly
to indicate that the Sun was born in a second- (10–1,000 K/h) compared to a cooling controlled
generation molecular cloud enriched in radioac- by free radiation into space. Because of these
tive elements by massive stars. At present, there is short timescales, one speaks of flash heating.
no satisfactory model accounting for the presence Most chondrules have been flash-heated several
of 26Al in our Solar System. It might result from an times, suggesting that a repetitive process was
improbable sequence of events, in which case the responsible for high-temperature processing. At
Solar System would be special in having hosted present, the most popular mechanism for chon-
26
Al while it formed. In such a case, our Solar drules formation is the shockwave model,
System might be unlike others in having evolved whereby a shockwave of speed 10 km/s impacts
(differentiated) parent bodies. Given that life, as dusty aggregates and heats them. The source of
we know it, is intimately linked to the geological these shockwaves in the protoplanetary disk is
history of our planet, it means that its development not well identified. Gravitational instabilities,
might result from a rare astrophysical event. X-ray flares, and planetesimals supersonic
motions have been proposed. All three models
High-Temperature Processes in the seem to face important difficulties.
Protoplanetary Disk
CAIs are very abundant in carbon-rich carbona- Formation and Early Evolution of Telluric
ceous chondrites, while they are virtually absent Planets
from other chondrites. They are especially large When the protoplanetary disk gas dissipated, we
and abundant in CV chondrites (▶ Meteorite, were left (in the inner Solar System) with a swarm
Allende). They therefore represent on the whole of planetesimals and planetary embryos. It took
a tiny fraction of chondritic matter. Understand- roughly 100 Ma to collect these bodies into the
ing their formation is, however, important terrestrial planets we know, such as the Earth.
because they are the first solids to have formed Growth occurred through random encounters of
in the protoplanetary disk. Their mineralogy is embryos. Though many embryos contributing to
compatible with that of condensation from a gas a given planet came from its neighborhood (the
of chondritic composition. Some of them show feeding zone), a significant number of embryos
evidence of remelting and evaporation. It is came from more distant regions.
widely believed that CAIs were formed close to Though accretion was mostly constructive,
the Sun (0.1 AU), where temperatures in the some impacts were also partly destructive. It is
disk were higher than 1,600 K. They were subse- widely believed that the Moon was made when a
quently transported to asteroidal distances, where planetary embryo, roughly the size of Mars, hit
chondrites formed, either by turbulent diffusion the proto-Earth. The Moon was built from that
of via magneto-hydrodynamic winds rooted at embryo, sometimes called Theia, and the Earth’s
the disk inner boundary. mantle. Thanks to the Hf-W isotopic system, it is
Chondrules are far more abundant than CAIs. possible to constrain that catastrophic event
They make up to 80 % of ordinary chondrites in which gave birth to our satellite. Latest measure-
volume. As ordinary chondrites represent more ments suggest it occurred 62þ90 10 Ma after Solar
than 80 % of the meteorites, chondrules were System formation, taken as the CAI formation.
574 Cosmochemistry
It is worth noting that dynamical studies dem- large craters seen today on the Moon date from
onstrated that the presence of the Moon stabilized that time. It is actually thanks to the return of the
the Earth’s obliquity. Were the Moon absent, the Moon rocks and their detailed study that the LHB
Earth’s obliquity would vary chaotically and was identified. If there was any life present on
Earth’s climate would have been far less stable Earth at that time, it may have been severely
than it has been. This singular, hazardous, event affected. Some other authors proposed that these
might therefore have played an important role in impacts could have, on the other hand,
the development of life as we know it. stimulated life.
While planets were forming, they developed a
magma ocean and iron cores appeared. Measure-
ments based on the Hf-W isotopic system dem- Future Directions
onstrated that the Martian core formation was
completed 10 Ma after the start of the Solar It is a challenging task to foresee the development
System. At present there is no undisputed esti- of a scientific discipline whose future depends on
mate for the age of the Earth’s core. the advancement of techniques, on the emergence
The origin of the terrestrial atmosphere is an of individuals, and on the general policy adopted
unsolved problem. It is a combination of delivery by a few prosperous countries which might not
of volatile compounds from extraterrestrial mat- remain so indefinitely. Some progress will be
ter and of a primordial, solar-type atmosphere made on the – already populous – data collection
trapped in the Earth’s mantle and subsequently front. That should serve to solve pending key
degassed. The excess of 129Xe (daughter isotope questions such as the homogeneity of short-
of 129I) in the Earth’s mantle compared to the lived radionuclides or the locus and timing of
Earth’s atmosphere indicates that the degassing organosynthesis. Instrumental developments
occurred early, possibly during the first 150 Ma. should also help to revisit dogma that is rarely
The origin of terrestrial oceans has long been disputed, such as the flash-heating model of
debated. Because they have a hydrogen isotopic chondrule formation for which evidence is thin.
composition different from that of Earth water, Laboratory experiments trying to reproduce
long-period comets such as comet Halley proba- key processes of the early Solar System should
bly did not contribute significantly to the water also be developed. Condensation and evaporation
budget. Jupiter-family comets and dark, water- experiments will illuminate the complex relation-
rich asteroids are the best candidates. Their D/H ships between gas and solids within the proto-
ratio – measured in carbonaceous chondrites – is planetary disk and establish chemical as well as
compatible with that of the terrestrial oceans. isotopic modifications induced by these funda-
These celestial bodies might also have delivered mental processes.
organic matter which might have provided pre- An almost virgin direction of research is the
cursors to life chemistry. The timing of Earth’ study of the mechanical property of meteorites.
water delivery relative to the formation of the Very little is known, for example, on their tensile
Moon is debated, though it seems pretty secure strength which is key to our understanding of
to state that water was delivered during the first planetary accretion and evolution. Such measure-
150 Ma. ments are desperately needed in the light of sev-
At that point, all bodies were formed, and the eral space missions.
architecture of the Solar System was only to be It is striking that, while a great deal is made of
significantly changed once, during the Late deuterium (2H) and 15N excesses in organic mat-
Heavy Bombardment (LHB) 3.8 Ga ago. At that ter, the processes that gave rise to them are still
epoch, after 0.8 Ga of slow migration, the giant debated. Developments on the modeling front are
planets Jupiter and Saturn crossed a resonance highly needed.
and destabilized a disk of planetesimals, which Cosmochemistry will certainly benefit from
were sent to the inner Solar System. Many of the increased interaction with astrophysics.
Cosmogonia: Greece 575
Observations of star-forming regions can be seen Brownlee DE et al (2006) Comet 81P/wild 2 under a
as a proxy of the environment of our Solar Sys- microscope. Science 314:1711–1716
Clayton DD, Nittler LR (2004) Astrophysics with presolar
tem formation. Thanks to space telescopes, such stardust. Annu Rev Astron Astrophys 42:39–78
as Herschel, or ground-based interferometers, Clayton RN, Grossman L et al (1973) A component of
such as ▶ ALMA, our understanding of accretion primitive nuclear composition in carbonaceous mete-
disks will be bettered, enabling us to “see” closer orites. Science 182:485–488
Engrand C, Maurette M (1998) Carbonaceous microme-
to the central star, in regions where precursors of teorites from Antarctica. Meteorit Planet Sci
C
planets similar to the Earth might form. Theoret- 33:565–580
ical astrophysics should be a daily companion of Gomes R, Levison HF, Tsiganis K, Morbidelli A (2005)
cosmochemists, and one could dream of a world Origin of the cataclysmic late heavy bombardment
period of the terrestrial planets. Nature 435:466–469
in which most, if not all, data collected are Krot AN, Scott ERD et al (2005) Chondrites and the
interpreted within a theoretical framework. protoplanetary disk. ASP conference series, San
One exigency could represent a horizon for Francisco
cosmochemistry: that of holding things together. Lauretta DS, McSween HY Jr (2008) Meteorites and early
solar system 2. Arizona University Press, Tucson
Too many disparate interpretations coexist without Lee T, Papanatassiou DA et al (1976) Demonstration of 26
being really coherent one with the other. Though Mg excess in Allende and evidence for 26Al. Geophys
some patches of the jigsaw puzzle have been thor- Res Lett 3:109–112
oughly assembled and though the global vision is Lodders K (2003) Solar system abundances and conden-
sation temperatures of the elements. Astrophys
probably correct, many of the data collected find J 591:1220–1247
no explanation within our current knowledge. In McCall GJH, Bowden AJ, Howarth RJ (2006) The history
that respect, astrophysics will, as always, teach us of meteoritics and key meteorite collections: fireballs,
humility. Though there is no reason for our Solar finds and falls. Geological Society Special Publica-
tions, London
System to be special, there are many possible out- Papike JJ (1998) Planetary materials. Mineralogical Soci-
comes of star and planetary formation. Ours is one ety of America, Washington, DC
among billions in the Galaxy. Patterson C (1956) Age of meteorites and the earth.
Shu FH, Shang H, Lee T (1996) Toward an astrophysical
theory of chondrites. Science 271:1545–1552
See Also
▶ Asteroid
▶ CAIs Cosmogonia: Greece
▶ Comet
▶ Geochronology Delphine Acolat
▶ Meteorite, Allende Centre François Viète, Université de Bretagne
▶ Meteorite, Murchison Occidentale, Quimper, France
▶ Meteorite, Orgueil
▶ Oxygen Isotopes
Keywords
References and Further Reading Universe; Greece; Myths; Origins; Earth;

Cosmos
Adams FC (2010) The birth environment of the solar
system. ARAA 48:47–85
Bernatowicz TJ, Fraundorf G, Ming T, Anders E,
Wopenka B, Zinner E, Fraundorf P (1987) Evidence Definition
for interstellar SiC in the Murray carbonaceous chon-
drite. Nature 330:728–730
Cosmogony is a theory about the origin and
Bogard DD, Nyquist L et al (1984) Noble gas contents of
shergottites and implications for the Martian origin of development of the cosmos or universe. The
SNC meteorites. GCA 48:1723–1739 word comes from the Greek kosmogonίa (from
576 Cosmogonia: Greece
kósmoB “cosmos,” “the world”) and the root of gί Gaia was the Protogenos (primeval divinity)
(g)nomai/gegona (“to be born, come about”). It is of earth, one of the primal elements who first
different from cosmology, which is the study of emerged at the dawn of creation, along with air,
the structure universe as a whole. Cosmos is not sea, and sky. She was the great mother of all
only a word for matter but for order, for process. palpable beings: the heavenly gods were
descended from her union with Ouranos (the
Sky), the sea gods from her union with Pontos
History (the Sea), the Gigantes from her mating with
Tartaros (the hell pit), and mortal creatures were
There are two types of Greek cosmogonies. One, sprung or born from her earthy flesh. With a
mythical, is very similar to the Mesopotamian cosmic order under the presence of Zeus, cos-
myths, based on divine couple who gives birth mogony is complete.
to all beings. The second, introduced by the But we also know the existence of variant
Greek philosophical tradition, is the first intellec- cosmogonies. In Homer’s Iliad (XIV, 210–246)
tual approach based on evidence, reason, and and later authors, Hera qualifies Oceanus and
debate. But the earliest speculations were still Tethys as the “father and mother of gods,” the
consistent with the mythical world structure. primordial couple. This can be establish an allu-
sion to a different cosmogony where primordial
chaos is watery.
Overview The Orphism (known by orphic fragments,
Greek hymns C3rd–C2nd BC), the religious
First Mythical Cosmogonies trend which took place away from the traditional
Hesiod’s Theogony (Yeogonίa), written in the practices of the cult, developed several cosmogo-
sixth century BC, is the first Greek mythical nies. It placed at the origin of the universe the
cosmogony: it is a large-scale synthesis of Night or the Time, which engenders a world egg,
Greek traditions concerning the gods. In the lan- primordial mixture of elements, which gives birth
guage of the Archaic period (eighth to sixth cen- to Gaia, Chaos, and Phanes, the primeval god of
tury BC), arche (a᾿ rwZ) designates the source, procreation, the driving force behind reproduction
origin, or root of things that exist. If a thing is to in the early cosmos. Phanes was the first king of
be well established or founded, its arche or static the universe, who passed the royal scepter on to his
point must be secure, and the most secure foun- daughter Nyx (Night), who in turn handed it down
dations are those provided by the gods: the inde- to her son Ouranos (Heaven). From him it was first
structible, immutable, and eternal ordering of seized by Khronos (Time) and then by Zeus, the
things. The initial state of the universe or the ultimate ruler of the cosmos. Phanes also incorpo-
origin (arche) is Chaos (wάoB): a gaping and rated the aspects of other primordial beings
indefinite void considered as a divine primordial described by various ancient writers including
condition from which everything else appeared. Thesis, Phusis, Ophion, Khronos, and Ananke.
Hesiod gives no idea about what preceded the All these gods are concepts rather than real
appearance of Chaos. characters.
Chaos was the first of the primordial deities,
followed by Gaia (Mother Earth), Tartaros (the Representations of Mother Earth and Phanes
underworld), the later-born Erebus (the darkness In Greek cosmology, Earth was conceived as a
between the earth and underworld), and Nyx flat disk encircled by the river Okeanos and
(Night). Eros is sexual desire – the urge to repro- topped above by the solid dome of heaven and
duce, not the emotion of love as is the common below by the great pit of Tartaros. She herself
misconception. Erebus and Nyx reproduced to supported the sea and mountains upon her breast.
make Aether (the outer atmosphere where the Mother Earth personification of the soil-
gods breathed) and Hemera (Day). ground whose products support human existence,
Cosmogonia: Greece 577
cosmogonical symbol of the material aspect of Anaximander of Miletus (610–545 BC)

the universe, was depicted as a buxom, matronly believed the beginning or ultimate reality is eter-
woman, half risen from the earth in Greek vase nal and infinite, or boundless (apeiron), subject to
painting and Roman mosaics (see neither old age nor decay, which perpetually
“▶ Cosmogonia: Roma”). She was portrayed as yields fresh materials from which everything we
inseparable from her native element. can perceive is derived.
Phanes was portrayed as a beautiful golden- Apeiron generated the opposites, hot-cold, C
winged hermaphroditic deity wrapped in a ser- wet-dry, etc., which acted on the creation of the
pent’s coils. The poets describe him as an incor- world. Everything is generated from apeiron and
poreal being invisible even through the eyes of then it is destroyed there according to necessity. It
the gods. His name means “bring to light” or acts as the substratum supporting opposites such
“make appear” from the Greek verbs phanaô as hot and cold and wet and dry and directed the
and phainô. movement of things, by which there grew up all
of the hosts of shapes and differences which are
Birth of Rational Cosmology found in the world. Out of the vague and limitless
There were many pre-philosophical cosmogonies body there sprang a central mass – this earth of
among the ancient Greeks, but the pre-Socratics ours – cylindrical in shape. A sphere of fire
and the Atomists differed fundamentally from all surrounded the air around the earth and had orig-
of them by seeking the origin of the world on a inally clung to it like the bark round a tree. When
rational basis rather than by appealing to a super- it broke, it created the sun, the moon, and the
natural force. stars.
Their understanding of the world applies the Anaximenes of Miletus (588–524 B.C.) posits
idea of order and harmony of a cosmos in all the air as the arche and ascribes to it divine attributes;
elements that have to obey physical and mathe- he provides a theory of change and supported it
matical laws. The greek science tries to report with observation. Using two contrary processes
most rationally the possibility of appearances of rarefaction and condensation (thinning or
observed in the sky, by elaborating explanatory thickening), he explains how air is part of a series
models which allow to foresee the return. of changes. Thinned air becomes fire and when
All of the pre-Socratics, only known by frag- condensed becomes first wind, then cloud, water,
ments, held that the cosmos has a beginning. earth, and stone in order.
Arche is basic stuff of the universe, the element For Heraclitus of Ephesus (540–480 BC),
and the first principle of existing things. This is arche is fire.
considered as a permanent substance or nature For Empedocles, it is composed of some com-
(physis) either one or more which is conserved bination of four elements: earth, water, fire, and
in the generation of the rest of it. From this all air. He is claiming that the primary opposites, hot
things first come to be and into this they are and cold and wet and dry, evolved from these
resolved in a final state. This source of entity is principles and that the other parts of the cosmos
always preserved (Aristotle, Metaph.A, evolved from these opposites.
983, b6ff). For the Atomists (Leucippus (fl. 430 BC) and
For Thales of Miletus (c. 624–547 BC), the Democritus (460–370 BC), the universe contains
arche is water. His theory was supported by the an infinite number of such indestructible beings,
observation of moisture throughout the world and “atoms”; space (as void) is not continuous, since
coincided with his theory that the earth floated on where atoms are, void is not.
water. His ideas were influenced by the Near- The generation of the gods is described by
Eastern mythological cosmogony and probably Plato (427–346 BC); but the origin of the cosmos
by the Homeric statement that the surrounding is philosophically explained by him, in the same
Oceanus (ocean) is the source of all springs and work, as the result of the planned actions of a
rivers. constructor or demiurge.
578 Cosmogonia: Mesopotamia
He conceived the earth as a sphere at the center different from cosmology, which is the study of
of the world. Aristotle taught that matter had nat- the structure universe as a whole.
ural motions depending on its composition.
The questions about the earth were summa-
rized by Aristotle (On the Heaven 1956, II 293a) History
with the question of the place of the earth in the
universe, of its shape, and of its size. In Mesopotamia the interest was more about ori-
gins and functions of gods than about the origins
of universe and life.
See Also “Water” was the prime element in all the tra-
ditions of Mesopotamia (Sumerians, Akkadians,
▶ Cosmogonia: Mesopotamia Hittites, Assyrians, etc.), and the roles of Tigris
▶ Cosmogonia: Roma and Euphrates were particularly central in myth-
ological texts.
Barbour IG (2006) Wissenschaft und Glaube. Historische Overview

und zeitgenössische Aspekte. Vandenhoeck &
Ruprecht, Göttingen In Mesopotamia, it seems at first glance that a
Curd P (1998) The legacy of Parmenides: eleatic monism much greater interest was taken in the origins and
and later presocratic thought. Princeton University
Press, Princeton functions of the gods than in the processes
Guthrie WKC (2000) A history of Greek philosophy. whereby the universe originated. The Mesopota-
Cambridge University Press, Cambridge mian myths and rituals do not actually deal with
Hansen W (2005) Classical mythology: a guide to the how and why the world was formed, but rather
mythical world of the Greeks and Romans. Oxford
University Press, New York how and why gods created specific things like
Kahn CH (1960) Anaximander and the origins of Greek human beings and living nature. In order to
cosmology. Columbia University Press, New York approach this ancient thought concerning the ori-
Kirk GS, Raven JE, Schofield M (2003) The presocratic gin of the cosmos, we have first to understand the
philosophers. Cambridge University Press, Cambridge
Rovelli C (2009) Anaximandre de Milet, ou la Naissance different cosmogonic associations between a
de la pensée scientifique. Dunod, Paris deity and prime elements of the universe. These
Sandwell B (1996) Presocratic philosophy, vol 3. associations may vary by regions and traditions
Routledge, New York over a period of more than 2,500 years of Meso-
Vernant JP (1964) Les origines de la pensée grecque. PUF,
Paris potamian history, within different cultures
Wright MR (1995) Cosmology in antiquity. Routledge, (Sumerians, Akkadians, Hittites, Assyrians, etc.).
London One of the main prime elements in these tra-
ditions is “Water.” The terms used in mytholog-
ical text are often Sea and River, which refer
Cosmogonia: Mesopotamia directly to the geographical description of Meso-
potamia with the main rivers, Euphrates and
Gregory Chambon Tigris, and the Arabic and Mediterranean seas.
Faculty of Arts and Humanities, University of The Babylonian creation epic “Enuma Elish,”
Nantes, Brest Cedex 3, France so-called from the first two words of the poem
“When on High” and probably composed in the
early part of the second millennium BC,
Definition describes how, during the very first separation
of order out of watery chaos, the primeval parents
Cosmogony is a narration about the origin and of the gods, Apsû (sweet waters underground)
development of the cosmos or universe. It is and Tiâmat (the sea), mingle their waters together
Cosmogonia: Roma 579
and create the second generation in the theogony. Lambert WG (1980–1983) Kosmogonie. In: Reallexikon
This second generation kills Apsû, who planned der AssyriologieVI. W. de Gruyter, Leipzig,
pp 218–222
to destroy his descendants, and asks Marduk, god Wiggermann F (1992) Mythological foundations of
of Babylon, for help and sent him against Tiâmat. nature. In: Diederick JW Meijer (ed.). Natural phe-
Armed with his winds and storms, he slices nomena. Their meaning, depiction and description in
Tiâmat’s body in half, forming Heaven with the the ancient Near East, Amsterdam, pp 279–306
upper part and Earth with the lower. Her mem- C
bers were used to supply its physical features: the
Tigris and Euphrates flow from her two eyes, the
bases of the Mountains were formed from her Cosmogonia: Roma
breasts, and her tail becomes the Milky Way.
The remainder of the story in the creation epic Delphine Acolat
deals with Marduk’s organization of the cosmos Centre François Viète, Université de Bretagne
with the offices that he assigns to the other gods Occidentale, Quimper, France
and his creation of man. The contrast between the
old chaos and the present order of the cosmos is
underlined by the difference of “dynamic” by the Keywords
gods: the second generation of the theogony is
considered as a generation of dynamic and mov- Universe; Roma; Myths; Origins; Earth; Cosmos
ing gods, whereas the chaos creatures Apsû and
Tiâmat stand for inertia and rest.
Others poems and epics show that Mesopota- Definition
mian tradition considered the physical universes
as a superimposition of levels. Above an under- Cosmogony is a theory about the origin and
world at the bottom, Apsû could be found as the development of the cosmos or universe. It is
dead body of the chaos monster of that name and different from cosmology, which is the study of
ruled by Ea, the god of wisdom, crafts, and crea- the structure universe as a whole.
tion. The second level consisted of the Earth,
where men were living and which was controlled
by Enlil, the god of breath and wind. The stars History
moved in an orbit above space, separating the
lower parts from their upper counterparts. Two Roman cosmogony is founded on those of earlier
heavens were situated at the top of these levels: classic age. The Romans took over Greek myths
the upper one was ruled by An, god of Heaven. and philosophic science (see Cosmogonia:
Greece) but added nothing to Greek cosmogony.
The Hellenistic view was relatively unchallenged
until the time of Nicolaus Copernicus
See Also (1473–1543) and Galileo Galilei (1564–1642).
▶ Cosmogonia: Greece
▶ Cosmogonia: Roma Overview
We can read Roman cosmogony by Ovid

References and Further Reading (43 BC–AD 17), Metamorphoses 1.1. He
described Chaos as “a rude and undeveloped
Bottéro J, Noah Kramer S (1992) Lorsque les Dieux
faisaient l’Homme. Gallimard
mass, that nothing made except a ponderous
Horowitz W (1988) The Babylonian map of the world. weight; and all discordant elements confused,
Iraq 50:147–165 were there congested in a shapeless heap” (see
580 Cosmogony
See Also
▶ Cosmogonia: Greece
▶ Cosmogonia: Mesopotamia
Barbour IG (2006) Wissenschaft und Glaube. Historische

und zeitgenössische Aspekte. Vandenhoeck &
Ruprecht, Göttingen
Bettini M (2011) Missing cosmogonies: the Roman case.
Archiv f€ur Religion geschichte, 13 http://antropologia-
mondoantico.files.wordpress.com/2013/08/bettini-
arch-rel-ge-2011-13.pdf. Accessed 10 Jun 2014
Hansen W (2005) Classical mythology: a guide to the
mythical world of the Greeks and Romans. Oxford
Cosmogonia: Roma, Fig. 1 Roman mosaic from a University Press, New York
Roman villa in Sentinum (now known as Sassoferrato, in Wright MR (1995) Cosmology in antiquity. Routledge,
Marche, Italy), ca. 200–250 A.D., Glyptotek Munich, Inv. London
W504). The Mother Earth goddess, Tellus is sitting with
four children, who represent the four seasons. Aion, the
god of eternity, is standing inside a celestial sphere deco-
rated with zodiac signs, in between a green tree and a bare
tree (summer and winter) Cosmogony
Daniel Rouan
Cosmogonia: Greece). Like ancient Greeks, LESIA, Observatoire Paris-Site de Meudon,
Romans begin the genesis of gods and nature Meudon, France
with a feminine entity, Gaia, which appears
after Chaos and before Eros. Chaos contains
primitive elements (earth, water, air, fire). Gaia Definition
is natural element which produces and supports
the world. In Ovid, an unknown god organizes all A cosmogony is any theory about the origin of the
these elements, and living beings occupy ether universe and/or the objects it contains. Most cul-
and stars (gods), air (birds), water (fishes), and tures developed their own cosmogony. In astron-
earth (animals and men). omy, the term usually refers to theories of
At the first century AD, Roman people pic- formation of the solar system.
tured the Mother Earth, supreme goddess of the
earth (Roman counterpart of Gaia) (Terra Mater/
Tellus), as a sweet mother: her role is reduced to
the dream giving and to the feeding of plants and COSPAR
children. Tellus thus refers to the guardian deity
of Earth and by extension the globe itself. The Catharine A. Conley
attributes of Tellus were the cornucopia or NASA Headquarters, Washington, DC, USA
bunches of flowers or fruit. She was typically
depicted reclining. In mosaic art, Gaia appears
as a full-figured, reclining woman, often clothed Definition
in green, and sometimes accompanied by grain
spirits – the Karpoi (e.g., in Antakya Museum, After the USSR launched its first Earth satellite in
inv. 847, the fourth century) (Fig. 1). 1957 and thereby opened the space age, the
Crater Chain 581
International Council of Scientific Unions

(ICSU), now the International Council for Sci- Counterglow
ence, established its Committee on Space
Research (COSPAR) during an international ▶ Gegenschein
meeting in London in 1958.
COSPAR’s objectives are to promote on an
international-level scientific research in space, C
with emphasis on the exchange of results, infor- Covalent Bonds
mation, and opinions, and to provide a forum
open to all scientists. These objectives are Henderson James (Jim) Cleaves II
achieved through the organization of Scientific Earth–Life Science Institute (ELSI), Tokyo
Assemblies, publications, and other means. Institute of Technology, Meguro–ku, Tokyo, Japan
COSPAR’s first Space Science Symposium was Institute for Advanced Study, Princeton, NJ, USA
organized in Nice in January 1960. Now general Blue Marble Space Institute of Science,
assemblies are organized every other year. Washington, DC, USA
In its first years of existence, COSPAR played Center for Chemical Evolution, Georgia Institute
an important role as an open bridge between East of Technology, Atlanta, GA, USA
and West for cooperation in space. When this role
became less prominent with the decline in rivalry
between the two blocs, COSPAR, as an interdis- Definition
ciplinary scientific organization, focused its
objectives on the progress of all kinds of research Covalent bonds are molecular bonds formed via
carried out with the use of space means the sharing of electrons according to Lewis rules to
(including balloons). form closed electron shells of two, four, or six
These activities are divided in “commissions,” electrons (for single, double, or triple bonds,
and inside the commission F is dedicated to life respectively). They are differentiated from ionic
sciences and subcommission F3 is devoted to and ▶ weak bonds by the amount of energy
astrobiology. required to break them, typically on the order of
COSPAR has also concerned itself with ques- 150–1,000 kJ/mole (1.5–10 eV/mole),
tions of biological contamination and spaceflight depending on the species. Covalent bonds tend to
while exploring the solar system. This proceeds form between atoms of similar electronegativity.
from Article IX of the Treaty on Principles Covalent bonds formed between atoms of dissim-
Governing the Activities of States in the Explo- ilar electronegativity (i.e., C-Cl) will be polar, with
ration and Use of Outer Space, Including the a greater amount of the electron density centered
Moon and Other Celestial Bodies (also known around the more electronegative nucleus.
as the UN Space Treaty of 1967). COSPAR,
based on the work of a dedicated planetary pro-
tection panel, proposes and maintains a planetary See Also
protection policy for the reference of spacefaring
nations. ▶ Hydrogen Bond
▶ Weak Bonds
See Also
▶ Cosmogonia: Greece Crater Chain

▶ Outer Space Treaty
▶ Planetary Protection ▶ Catena, Catenae
582 Crater Lakes (Mars)
See Also
Crater Lakes (Mars)
▶ Crater, Impact
Alessandro Airo ▶ Habitability on Mars
Institut f€
ur Geologische Wissenschaften
Tektonik und Sedimentäre Geologie, Freie
References
Geowissenschaften, Berlin, Germany McKay CP, Davis WL (1991) Duration of liquid water
habitats on early Mars. Icarus 90:214–221
Michalski JR, Cuadros J, Niles PB, Parnell J, Rogers AD,
Wright SP (2013) Groundwater activity on Mars and
implications for a deep biosphere. Nat Geosci
Definition 6:133–138
Mouginis-Mark PJ, Boyce JM (2012) Tooting crater:
It is widely accepted that liquid water was present geology and geomorphology of the archetype large,
on the surface of ▶ Mars during its early history. fresh, impact crater on Mars. Chem Erde 72:1–23
Pondrelli M, Baliva A, Di Lorenzo S, Marinangeli L,
Consequently, this surface water must have Rossi AP (2005) Complex evolution of paleolacustrine
migrated downhill, accumulating in topographic systems on Mars: an example from the Holden crater.
depressions. Besides the ocean-size northern J Geophys Res 110:1–20
lowlands, the most prominent basins on Mars
are the numerous impact craters. There is much
evidence for prolonged water ponding within
individual or interconnected impact craters, Crater, Impact
such as (1) channels leading into the craters and
having deposited delta fans, (2) the presence of Roland J. Wagner
paleoshorelines along the crater walls, or (3) the German Aerospace Center (DLR), Institute of
occurrence of horizontally layered deposits com- Planetary Research, Berlin, Germany
posed of phyllosilicates or sulfates interpreted to
represent lacustrine sediments and evaporites
(e.g., Michalski et al. 2013; Pondrelli et al. 2005). Keywords
Although there are no substantial amounts of
liquid water on Mars’ surface today, a sufficiently Asteroid; Collision; Comet; Impact; Micromete-
large impact into ground-ice baring terrain could orite; Surface
potentially create a transitional crater lake.
Impact-induced ground-ice melting is supported
by the presence of fluidized ejecta around ram- Definition
part craters, which are thought to require liquid
water for their formation (Mouginis-Mark and An impact crater is a mostly circular or elliptical
Boyce 2012). Under current Mars conditions, a elongate depression, generally with a raised rim,
lake would quickly freeze at the surface, but which is created by the impact of a minor body on
modeling studies predict that such an ice layer the solid surface of a ▶ planet or satellite. Impact
would protect the underlying water from rapid craters range in size from microcraters seen only
evaporation and therefore preserve the lake for microscopically on surfaces of ▶ rock samples
up to a few hundreds of thousands of years (e.g., rocks from the lunar surface) to large craters
(McKay and Davis 1991). Furthermore, it can and ▶ impact basins several hundreds or thou-
be assumed that the impact-related heating of sands of kilometers across. Impacts into the
the crater floor would delay the freezing of the atmospheres of the large gaseous planets in the
entire lake water and potentially drive long-term outer ▶ Solar System can produce transitory cir-
hydrothermal circulation. cular or semicircular features resembling craters,
Crater, Impact 583
as has been observed on ▶ Jupiter after the include (Pike 1980; Chapman and McKinnon
impact of ▶ Comet Shoemaker/Levy-9. 1986; Melosh 1989; Schenk et al. 2004) (a) flat
crater floors, (b) terraces at crater wall interiors,
(c) central peaks, or (d) peak rings. On icy satel-
Overview lites, complex crater forms include (e) central pits
(also observed on Mars [e.g., Barlow 2009]),
Impact craters on solid surfaces of planets and (f) central domes, or (g) bright, almost flat circu- C
satellites are created by hypervelocity collisions lar areas termed palimpsests (Faculae) devoid of
with smaller bodies. These bodies, termed impac- prominent topographic features such as
tors or projectiles, range in size from ▶ micro- crater rims.
meteorites to large bolides up to tens or hundreds The largest impact structures are ▶ impact
of kilometers in diameter, which form impact basins, which exhibit two or even more rings
basins several hundreds or thousands of kilome- (ridges or graben) and are termed multi-ring
ters across (Pike 1980; Melosh 1989). Candidate basins (e.g., Spudis 1993).
impactors are (1) ▶ asteroids from the main belt
(MBA) or from other asteroid families, for exam-
ple, Near-Earth asteroids (NEA) (Neukum See Also
et al. 2001; Strom et al. 2005; Bottke
et al. 2012), (2) ▶ comets including ▶ ecliptic ▶ Asteroid
or short-period comets (EC, orbital period ▶ Asteroid Belt, Main
<200 years) derived from the ▶ Kuiper belt and ▶ Catena, Catenae
nearly isotropic or long-period comets (NIC, ▶ Chronostratigraphy
orbital period >200 years) from the ▶ Oort ▶ Comet
cloud (Zahnle et al. 2003), (3) bodies or debris ▶ Comet Shoemaker-Levy 9
in planetocentric orbits (Neukum 1985; Chapman ▶ Ecliptic
and McKinnon 1986), and (4) remnants of plan- ▶ Facula, Faculae
etary accretion (planetesimals) (Wetherill 1975). ▶ Impact Basin
The number or frequency of craters on a sur- ▶ Jupiter
face per unit area records its age: the higher the ▶ Kuiper Belt
crater frequency, the older the age of the surface ▶ Mars
due to the longer exposure time to the incoming ▶ Micrometeorites
impactor flux. This relationship can be used as an ▶ Oort Cloud
important tool in planetary chronology. ▶ Planet
Morphology and sizes of impact craters reflect ▶ Planet Formation
impact conditions, projectile properties, target ▶ Planetesimals
properties, and changes of target properties with ▶ Rock
time (Schenk et al. 2004). The smallest craters ▶ Satellite or Moon
identified in camera images are simple craters, ▶ System Solar Formation, Chronology of
characterized by a bowl-shaped, parabolic crater
morphology (e.g., Melosh 1989). With increasing
diameter, crater forms become more complex.
The simple-to-complex transition diameter Barlow NG (2009) Martian central pit craters: summary of
approximately scales with the inverse of the grav- northern hemisphere results. Lunar and planetary sci-
ity acceleration, except for icy surfaces as on ence conference 40th, abstr. 1915 [CD-Rom]
▶ Mars (ice in the regolith) and the icy satellites Bottke WF, Vokrouhlicky D, Minton D, Nesvorny D,
Morbidelli A, Brasser R, Simonson B, Levison HF
in the Outer Solar System (Chapman and (2012) An Archean heavy bombardment from a
McKinnon 1986; Melosh 1989; Schenk destabilized extension of the asteroid belt. Nature.
et al. 2004). Features observed in complex craters doi:10.1038/nature10967
584 Cratering Chronology
Chapman CR, McKinnon WB (1986) Cratering of plane- typically composed of granitoid, high-grade,
tary satellites. In: Burns JA, Matthews MS (eds) Sat- strongly deformed metamorphic rocks and
ellites. University of Arizona Press, Tucson,
pp 492–580 less metamorphosed metavolcanic and
Melosh HJ (1989) Impact cratering: a geologic process, metasedimentary rocks (▶ greenstone belts).
vol 11, Oxford monographs on geology & geophysics. The crustal section of a craton is thicker than
Oxford University Press, New York, p 245 that of normal continental crust (which is
Neukum G (1985) Cratering records of the satellites of
Jupiter and Saturn. Adv Space Sci 5:107–116 ~30–40 km) and is underlain by a thick (up to
Neukum G, Ivanov BA, Hartmann WK (2001) Cratering several hundred km) root of low-density,
records in the inner solar system in relation to the lunar depleted lithospheric mantle. Cratons formed
reference system. In: Hartmann WK, Geiss J, through orogenies (mountain-building pro-
Kallenbach R (eds) Chronology and evolution of
Mars. Kluwer, Dordrecht, pp 53–86 cesses) and accretion of crustal fragments dur-
Pike RJ (1980) Control of crater morphology by gravity ing the Archean when mantle and crustal
and target type: Mars, Earth, Moon. In: Proceedings of temperatures were higher than those of today.
the lunar and planetary science conference 11th, Hous- When covered by younger sedimentary basins,
ton, pp 2159–2189
Schenk PM, Chapman CR, Zahnle K, Moore JM they are referred to as platforms. Examples of
(2004) Ages and interiors: the cratering record of the well-known Archean cratons are the Slave Cra-
Galilean satellites. In: Bagenal F, Dowling T, ton of northwest Canada, the ▶ Pilbara Craton of
McKinnon W (eds) Jupiter – the planet, satellites and northwest Australia, and the Kaapvaal Craton of
magnetosphere. Cambridge University Press, Cam-
bridge, pp 427–456 southern Africa.
Spudis PD (1993) The geology of impact basins: the moon
and other planets. Cambridge University Press, Cam-
bridge, 263 pp
Strom RG, Malhotra R, Takashi I, Yoshida F, Kring DA
(2005) The origin of planetary impactors in the inner See Also
solar system. Science 309:1847–1850
Wetherill GW (1975) Late heavy bombardment of the
moon and the terrestrial planets. Proceedings of the ▶ Archean Eon
lunar and planetary science conference 6th, Houston, ▶ Continental Crust
pp 1539–1561 ▶ Crust
Zahnle K, Schenk P, Levison H, Dones L (2003) Cratering ▶ Granite
rates in the outer solar system. Icarus 163:263–289
▶ Shield
Cratering Chronology
▶ Chronostratigraphy
Crenarchaeota
Craton Ricardo Amils

Nicholas Arndt Universidad Autónoma de Madrid, Madrid,
ISTerre, Université Grenoble Alpes, France Spain
A craton is an old, thick, cool, and rigid part of Crenarchaeota is one of the four phyla of
the continental ▶ lithosphere. Its crustal part is ▶ Archaea. Crenarchaeota comprises both
Crust 585
hyperthermophilic and cold-dwelling prokary-

otes. The hyperthermophilic species of Critical Core Mass (Giant Planet
Crenarchaeota tend to cluster closely together Formation)
and occupy short branches on the 16S rRNA
gene phylogenetic tree. These organisms are con- Yann Alibert1 and Ravit Helled2
1
sidered good models for early Archaea. Phyloge- Space Research and Planetary Sciences, Physics
netically, these are a more rapidly evolving Institute, University of Bern, Bern, Swiss C
2
species. Most hyperthermophilic Crenarchaeota Geophysical, Atmospheric and Planetary
have been isolated from geothermal heated soils, Sciences, Tel Aviv University, Raymond and
waters containing elemental sulfur and sulfides, Beverly Sackler Faculty of Exact Scienes, Tel
or hydrothermal vents. Among the hyperthermo- Aviv, Israel
philic Crenarchaeota, we can find members of the
Sulfolobales, Thermoproteales, and Desulfuro-
coccales orders. One species of Sulfolobus, Keywords
S. acidocaldarius, was the first hyperthermo-
philic Archaea discovered. It was isolated by Planet Formation; Core accretion (Model for
Thomas Brock and colleagues in Yellowstone Giant Planet Formation)
National Park, U.S.A., in 1970. Special mention
should be given to Pyrodyctum fumarii that can Definition
grow at 113 C.
In the Core Accretion model for giant planet for-
mation, the critical core mass is the minimum core
mass that is required to initiate rapid gas accretion,
which leads to the formation of a gaseous planet.
See Also
▶ Acidophile See Also

▶ Archaea
▶ Core Accretion, Model for Giant Planet
▶ Hyperthermophile Formation
▶ Phylogeny
▶ Sulfur Cycle
▶ Yellowstone National Park, Natural Analogue
Site
Crossing Over
▶ Recombination
Cretaceous/Paleogene (KPg)
Crust
▶ KT Boundary
Nicholas Arndt
Definition
Cretaceous/Tertiary Boundary (KT)
The crust is the outer rocky layer of the Earth. It
▶ KT Boundary also refers to the outer rocky layer of telluric
586 Crustal Deformation
planets or moons. The Earth’s crust has a rela-

tively low density and floats on the underlying Cryosphere
mantle. ▶ Oceanic crust, which covers about two
thirds of the Earth’s surface, is 6–9-km thick and Nicholas Arndt
composed mainly of basalt. ▶ Continental crust ISTerre, Université Grenoble Alpes, France
is thicker (about 10-km thick in rifted portions,
>80-km thick beneath active mountain belts),
averaging about 30 km. Continental crust is com- Definition
posed mainly of granitic and metamorphic rocks.
Oceanic crust forms at spreading centers and is The cryosphere is that part of the Earth and other
never older than 200 Ma. Continental crust has planets in which temperatures are low and water
formed continuously throughout the Earth’s his- is solid. On Earth, it constitutes the polar ice caps
tory, from 3.8 or even 4.3 Ga. Mars and probably and pack ice, mountain glaciers, cold deserts, and
Venus have basaltic crusts. The lunar crust is regions of permafrost. During the time of near-
basaltic in maria and composed of anorthositic global ▶ glaciations in the Neoproterozoic – the
breccia in lunar highlands. Cryogenian famous for the “▶ Snowball Earth”
hypothesis – the cryosphere encompassed almost
all the planet, as is the case at present time for icy
See Also moons such as ▶ Europa.
▶ Anorthosite
▶ Continental Crust See Also
▶ Craton
▶ Crust ▶ Europa
▶ Kreep ▶ Glaciation
▶ Moon, The ▶ Snowball Earth
▶ Oceanic Crust
▶ Plate Tectonics
Cryostat
Crustal Deformation
Inge Loes ten Kate
▶ Archean Tectonics Earth Sciences, Utrecht University, Utrecht,
The Netherlands
Cryocooler Synonyms
▶ Cryostat Cryocooler; Dewar flask
Definition
Cryophile
A cryostat is an apparatus used to maintain
▶ Psychrophile very low (“cryogenic” <100 K) temperatures.
Cryptoendolithic 587
It typically consists of two vessels, one mounted See Also

inside of the other. The inner vessel contains
the cold sample (cryogen) mounted inside an ▶ Enceladus
evacuated outer vessel. The vessels are held ▶ Io
together by a material with low thermal conduc- ▶ Titan
tivity. The vacuum in the outer vessel serves as ▶ Trans-neptunian Object
a thermal insulator. The two vessels are C
separated by a radiation shield to prevent heat
transfer. The radiation shield is cooled by a
cryocooler.
In medicine, a cryostat is a device to cut Cryptoendolithic
histological slides, consisting of a microtome
(ultrathin slicer) in a freezer. In astrobiology, Felipe Gomez
cryostats are used to generate low temperature Centro de Astrobiologı́a (CSIC/INTA), Instituto
conditions, for example, for simulating surfaces Nacional de Técnica Aeroespacial, Torrejón de
of icy moons or Titan or even the surface of Ardoz, Madrid, Spain
Mars. Furthermore, studies of ices and chemis-
try taking place in ices require the use of a
cryostat; examples of these are interstellar and Definition
cometary ices.
Cryptoendolithic refers to one of the three sub-
classes in which ▶ endolithic microorganisms are
classified. Cryptoendolithic microorganisms are
those able to colonize the empty spaces or pores
Cryovolcanism inside a rock with the connotation of being hid-
den. This connotation is important to astrobiol-
Therese Encrenaz ogy as these protected environments inside rocks
LESIA, Observatoire de Paris - Section de are putative habitable niches where life could be
Meudon, Meudon, France sustained in very adverse conditions or space
environments. Cryptoendolithic microorganisms
can survive on inorganic metabolites from the
Definition surroundings; thus, these microbes are mainly
lithotrophs. Depending on the physicochemical
Cryovolcanism is a phenomenon that occurs in properties of the mineral structure of the rock, it
environments with extremely low temperature can provide protection against damaging radia-
and resembles silicate volcanism. There, instead tion. Some reported examples have been
of molten silicates, cryovolcanoes erupt liquid described in basaltic rocks from Antarctica
water, methane, and ammonia onto the icy sur- where low temperatures are very restrictive for
face of a planet or satellite. It has been observed life or from high-altitude environments with high
on several satellites of the outer Solar System. In radiation doses.
particular, active cryovolcanism has been discov-
ered on Enceladus, one of Saturn’s satellites.
Traces of cryovolcanism are also found on See Also
▶ Titan and ▶ Triton. Cryovolcanism could also
be present on other outer satellites and trans- ▶ Chemolithotroph
Neptunian objects. ▶ Endolithic
588 CS
also several sites that could be studied or used

CS as Martian analogues.
Canada is pursuing a strategy to develop its
▶ Carbon Monosulfide expertise in remote sensing, space robotics, and
space telecommunication.
In 2010, the CSA employed around 700 full-
time equivalents.
CSA
Michel Viso 1934 CT

CNES/DSP/SME, Vétérinaire/DVM, Astro/
Exobiology, Paris Cedex 1, France ▶ Toutatis
Definition
Culture Media
The Canadian Space Agency was established in
1989 and is committed to lead the development ▶ Macronutrient
and application of space knowledge for the ben-
efit of Canadians and humanity. Canada began
space activities at the early stage of the space era,
through an agreement with the US National Aero- Curiosity
nautic and Space Administration (▶ NASA) to
build and launch satellites to study the upper ▶ Mars Science Laboratory
atmosphere. For several years from 1958, Canada
operated jointly with the USA, the Fort Churchill
base in Manitoba, to launch sounding rockets. In
1969, the federal government created Telesat Cuvier’s Conception of Origins of Life
Canada to build and exploit Canadian communi-
cation satellites. The government also set up a Stéphane Tirard
Department of Communications that immedi- Centre François Viète d’Histoire des Sciences
ately took over from the Communications et des Techniques EA 1161, Faculté des
Research Centre and the Interdepartmental Com- Sciences et des Techniques de Nantes, Nantes,
mittee on Space. In 1974, NASA awarded Can- France
ada the responsibility of designing, developing,
and building the Shuttle Remote Manipulator
System (SRMS) for the Space Shuttle. This History
agreement resulted in Canadarm, the shuttle’s
15-m robotic arm, and led to the flight, in 1984, The French naturalist Georges Cuvier
of the first Canadian astronaut (Marc Garneau). (1769–1832) was one of the most important com-
In 1979, Canada became an associate member of parative anatomists and paleontologists of the
the European Space Agency (ESA), and in 1985, beginning of the nineteenth century. Concerning
Canada accepted to participate in the ▶ Interna- history of life, Cuvier claimed a form of fixism
tional Space Station program. Canadian scientists explaining changes of species during geological
are cooperating worldwide in manned space time. He imagined several disasters during which
flight, space sciences, and exploration of the certain species would disappear and after which
solar system. For astrobiology, Canada offers new species would come from other places.
Cyanoacetylene 589
During the first part of the nineteenth century, his H2 NCN þ NH3 ! H2 NCðNÞNH2 (2)
proposal had a very significant place in biology
and paleontology. Reaction with amines, such as amino acids, gives
N-carbamoylamino acids and hydantoins.
See Also
See Also C
▶ Darwin’s Conception of the Origins of Life
▶ Lamarck’s Conception of Origins of Life ▶ Hydantoin
Cyanamide Cyanic Acid
Henderson James (Jim) Cleaves II ▶ HCNO Isomers

Earth–Life Science Institute (ELSI),
Tokyo Institute of Technology, Meguro–ku,
Tokyo, Japan
Institute for Advanced Study, Princeton, Cyanide Anion
NJ, USA
Blue Marble Space Institute of Science, ▶ CN-
Washington, DC, USA
Cyanide Ion
Synonyms ▶ CN-
Amidocyanogen; Carbamonitrile; Carbimide;

Carbodiimide; Cyanoamine; Cyanogenamide;
Cyanogen nitride; H2NCN; Hydrogen cyana- Cyano Radical
mide; N-Cyanoamine
▶ Cyanogen Radical
Definition
Cyanamide is a simple compound (H2NCN) Cyanoacetylene

formed by the irradiation of cyanide. Cyanamide
has been detected in the ▶ interstellar medium José Cernicharo Quintanilla
(see ▶ Molecules in Space) and has been shown Department of Astrophysics, Laboratory of
to be an effective condensation agent for both Molecular Astrophysics, Iorrejón de Ardoz,
peptides and nucleotides. Reaction of cyanamide Madrid, Spain
with water yields urea:
H2 NCN þ H2 O ! H2 NCONH2 (1) Synonyms
Reaction with ammonia gives guanidine: CA

590 Cyanoamine
Definition
Cyanobacteria
Cyanoacetylene, HC3N (H–CC–CN), is an
organic molecule that is the simplest Josef Elster1,2 and Jana Kvı́derová2
1
cyanopolyyne (H(–CC–)n–CN). It was one Faculty of Science, Centre for Polar Ecology,
of the first molecules detected in space using University of South Bohemia, Ceske Budejovice,
radio astronomical techniques (Turner 1971). Czech Republic
2
The observation of its rotational lines is used in Institute of Botany, Academy of Sciences of the
radio astronomy to derive the physical condi- Czech Republic, Trebon, Czech Republic
tions of interstellar and circumstellar clouds. It
is also an important component of Titan’s atmo-
sphere (Kunde et al. 1981), where it is found in Keywords
the gas phase in the upper atmosphere and in ice
form in the lower stratosphere (see, e.g., Ander- Antarctic; Carbon and nitrogen cycles; Desicca-
son et al. 2010). It is a trace constituent in com- tion; Endosymbiosis; Extremophiles; Freeze-
etary atmospheres (comae). Cyanoacetylene has melt stress; Halophily; Irradiance; Limits of sur-
been proposed as a prebiotic reagent for the vival; Oxygenic photosynthesis; Psychrophily;
formation of pyrimidine bases, nucleosides, Thermophily; Ultraviolet radiation
and nucleotides (Sanchez and Orgel 1970;
Powner et al. 2009).
Synonyms
See Also Blue-green bacteria; Cyanophyceae
▶ Titan
Definition
References and Further Reading ▶ Cyanobacteria are photosynthetic ▶ bacteria

that use ▶ water as reducing power to release O2.
Anderson CM, Samuelson RE, Bjoraker GL, Achterberg They evolved early in Earth’s history. As bacterial
RK (2010) Particle size and abundance of HC3N ice in
Titan’s lower stratosphere at high northern latitudes.
primary producers, cyanobacteria occupy a
Icarus 207:914 privileged position among organisms due to their
Kunde VG, Aikin AC, Hanel RA, Jennings DE, Maguire role in the carbon and nitrogen cycles. They are
WC, Samuelson RE (1981) C4H2, HC3N and C2N2 in widely adapted to different extreme environments
Titan’s atmosphere. Nature 292:686
and play an important role, especially in cold polar
Powner MW, Gerland B, Sutherland JD (2009) Synthesis
of activated pyrimidine ribonucleotides in and alpine environments, because of their toler-
prebiotically plausible conditions. Nature 459:239 ance of a wide temperature range, ▶ desiccation,
Sanchez RA, Orgel LE (1970) Studies in prebiotic syn- freeze-melt, and salinity stress.
thesis. V. Synthesis and photoanomerization of pyrim-
idine nucleosides. J Mol Biol 47:531
Turner BE (1971) Detection of interstellar
cyanoacetylene. Astrophys J 163:L35 Overview
Cyanobacteria are oxygenic photosynthetic pro-

karyotes responsible for the transformation of a
reduced atmosphere to an oxidized one. The oxy-
Cyanoamine gen produced by this photosynthetic group of bac-
teria drove life to adapt to the newly formed
▶ Cyanamide aerobic environments, resulting in the evolution
Cyanobacteria 591
of novel physiologies, biochemistries, and mor- Many cyanobacteria can develop in extreme
phologies. In addition, cyanobacteria, as the pho- environments, at extremely high (geothermal
tosynthetic partner in the primary endosymbiotic springs) and low (Antarctic, Arctic, alpine areas,
event, introduced photoautotrophy to eukaryotes. permafrost) temperatures, in hypersaline
The geopaleological record indicates that (halophiles) and alkaline (alkaliphiles) habitats,
cyanobacterial ▶ photosynthesis took place early under high radiation conditions, desiccation, and
in our planet’s history. As primary producers, toxicity stress, etc. A brief survey follows the C
cyanobacteria occupy a privileged position occurrence of cyanobacteria in different extreme
among organisms because of their role in carbon environments.
and nitrogen (N-fixers) biological cycles (Whitton
and Potts 2000; Knoll 2008; Swingley et al. 2008). Thermophilic Cyanobacteria
In comparison with other groups of bacteria, Results of recent molecular biological studies
cyanobacteria exhibit an unusually wide range of provide new information about cyanobacteria
morphologies. Traditional taxonomic that inhabit high-temperature habitats.
cyanobacterial classification is based on morphol- A number of earlier reports proposed that ther-
ogy and development, recognizing five principal mophilic cyanobacteria had branched out at the
groups: Group I, Chroococcales, solitary and colo- beginning of their evolution. However, recent
nial unicellular forms; Group II, Pleurocapsales, studies have brought arguments for a later emer-
unicellular to pseudo-filamentous, with cells capa- gence of thermophilic cyanobacteria. The present
ble of multiple and binary fission; Group III, analyses (Ward and Castenholz 2000) strongly
Oscillatoriales, filamentous forms with ▶ cell dif- confirm two distinct thermophilic lineages. Geo-
ferentiation; Group IV, Nostocales, filamentous thermal springs can be considered as isolated
with cell differentiation to produce akinetes and islands and therefore an ideal site for the origin
heterocysts; and finally Group V, Stigonematales, and evolution of endemic species. Some species
with cell differentiation and complex multicellular of Thermosynechococcus spp. are clearly
organization (Komárek and Anagnostidis 1998, restricted in geographical distribution, although
2005). Results of the application of recent molec- other thermophilic cyanobacteria, such as
ular techniques (e.g., sequence of the 16 S rRNA Mastigocladus laminosus and Cyanothece
gene) support some but not all these groups. Fea- minervae, appear to be cosmopolitan.
tures like cell differentiation or multiple fissions Cyanobacteria occupying geothermal springs
are supported by molecular analyses, whereas uni- are not observed below ▶ pH 4, and their diver-
cellular forms and simple filaments do not gener- sity is quite restricted at pH below 6. Tempera-
ate monophyletic groups. It has been possible to ture, in combination with availability of nitrogen
reconstruct an evolutionary history of the and the presence of free sulfide, determines the
cyanobacterial groups, establishing a framework cyanobacterial species composition, because sul-
for resolving how their metabolic and phenotypic fide is an efficient inhibitor of oxygenic photo-
diversity came about (Six et al. 2007). synthesis. The detected upper-temperature limit
Probably due to their evolutionary antiquity, for cyanobacteria (Synechococcus lividus) and
cyanobacteria are widely adapted to all extremes for global photosynthesis is presently 73–74 C.
related to changes in geological time (Elster
et al. 2001). Tolerance of low-oxygen conditions Cyanobacteria Under Low Temperature,
is still widespread among cyanobacteria and free Desiccation, and Salinity Stress
sulfide is tolerated by some strains. In addition, Cyanobacteria, frequently considered warm-
some cyanobacterial strains can use H2S as a water organisms, play an important role in the
hydrogen donor. They also tolerate high doses carbon and nitrogen cycles in cold polar and
of ultraviolet B and C radiation. All these features alpine environments. The ecophysiological fea-
have been especially important in the early evo- tures that predetermine their dominance in these
lution of cyanobacteria. environments include: slow growth rate over a
592 Cyanobacteria
wide temperature range, tolerance to desiccation, revealed the similarity of strains from Ellesmere
freeze-melt and salinity stress, and a variety of Island, the Canadian Arctic, and Abisko, Sweden,
photoacclimation strategies to both high and low with strains from Svalbard. The rate of biological
solar irradiance. In cold environments, where colonization of the Arctic from southern latitudes
invertebrate grazing pressure is limited, the is relatively high (Strunecký et al. 2012b).
large standing stocks of cyanobacterial biomass Most polar and alpine cyanobacterial species
are the result of its gradual accumulation over tested up to now are rather more psychrotolerant
many seasons (Vincent 2000). than psychrophilic. Polar cyanobacteria have long
At present, endemism of polar and alpine doubling times in comparison with psychrophilic
cyanobacteria is under discussion. Castenholz eukaryotic ▶ algae and heterotrophic bacteria.
(1992) noted that due to the slow rates of speci- Diverse evidence indicates that the
ation and high dispersal abilities together with the unsaturation of ▶ membrane lipids correlates
relatively young age of most polar ice-free areas, with low-temperature sensitivity, although this is
cyanobacterial endemism in extremely cold envi- not the only factor that regulates low-temperature
ronment is improbable. However, recent studies resistance in cyanobacteria. The unsaturation
based on 16S rDNA and 16S–23S rDNA apparently protects the photosystem II complex
intergenic spacer sequences from the Antarctic from low-temperature photoinhibition.
and Arctic (Taton et al. 2006; Comte et al. 2007; Cyanobacteria have several strategies to min-
Komárek et al. 2008; Strunecký et al. 2010, imize osmotic and mechanical stresses. They are
2012a, b) introduced that for selected able to produce mucopolysaccharides
cyanobacterial clusters apparently belonging to (exopolymeric substances), which slow down
generic entities, geographical limitation plays a the flow of liquid water during freeze up and
prominent role. The north and south polar regions thaw (Vincent 2000). Cyanobacteria also respond
are remarkably different from each other in their to these stresses by producing compatible solutes.
geological histories, biodiversity, energy bal- Large disaccharide sugars, such as trehalose,
ance, and transport, yet they share the common sucrose, and glucosylglycerol, are typical com-
challenges for the life that exists there, the fluc- patible solutes in water-stressed cyanobacteria
tuating extremes of cold, and desiccation stress. (Reed et al. 1984). The extreme tolerance of
The Arctic is a geologically young and open area cyanobacteria to desiccation is exemplified by
in which rapid glacial and periglacial processes their ability to tolerate very low water potential.
occur. There is a relatively (to Antarctica) higher Chroococcus and Chroococcidiopsis can both fix
level of biodiversity and a greater capacity for CO2 at remarkably low water potentials.
geographical genome transfers to take place, Cyanobacteria are an important component of
aided by biological vectors, land mass continuity, hypersaline ecosystems. Compatible solutes are
and air and oceanic currents. In contrast, a thick especially vital for cyanobacterial survival in
ice sheet covers most of the Antarctic continent, saline desert evaporate soils when they are
which has been geographically isolated for a very under the combined stress of desiccation and
long period of time. As a consequence, glacial hypersaline conditions (Oren 2000). In these hab-
and periglacial processes are much slower, and itats, glycine betaine is the most common com-
genome exchange and movement, and conse- patible solute. A wide range of species belonging
quentially biodiversity, are more limited in the to different taxonomical groups has been reported
south polar region of Antarctica (Elster and Ben- to thrive at high salt concentrations (e.g.,
son 2004). It has been tested that cyanobacteria Microcoleus chthonoplastes, Oscillatoria
survived Antarctic glaciations directly on site limnetica, Synechocystis spp.).
after the Gondwana breakup by using the relaxed
and strict molecular clock in the analysis of the Light and UV Radiation
16S rRNA gene (Strunecký et al. 2012a). In the Under natural conditions, cyanobacteria experi-
contrary, in the Arctic, molecular studies ence light conditions that fluctuate rapidly,
Cyanobacteria 593
frequently reaching suboptimal levels for photo- ▶ microorganisms including photosynthetic

synthesis. Both the intensity and quality of the cyanobacteria to be determined. Since
irradiance can vary dramatically during the day cyanobacteria dominate in the polar regions, they
and among habitats. In desert terrestrial environ- can serve as model organisms for evaluation of
ments, irradiance may be very bright and cause survival at low temperatures and other stress factors
photoinhibition or damage to the reaction centers common to the polar ▶ environment. In the most
of PSII (photooxidative bleaching). extreme Antarctic conditions, cyanobacteria play a C
Cyanobacteria can modify the protein composi- determining role in (1) desert communities, includ-
tion of PSII at high irradiance, making the PSII ing soil crust and ▶ epilithic and ▶ endolithic spe-
reaction centers less susceptible to cies of cold polar deserts, e.g., in Dry Valleys in the
photoinhibition. Rapid light intensity or quality Antarctic, where the conditions resemble those of
fluctuations result in rapid photoacclimation pro- the ▶ Mars surface (Friedmann 1982), and (2) ben-
cesses, such as state transition that can modify the thos of permanently frozen lakes, e.g., Lakes Hoare
photosynthetic apparatus within minutes. These and Fryxell in the Antarctic, which represent
short-term modifications usually do not need pro- another environment that could occur on Mars
tein synthesis. For example, during the state tran- (de Pablo et al. 2008). The conditions of perma-
sitions, the phycobilin antennae migrate between nently ice-covered lakes and seas could also be
the photosystems to optimize the distribution of relevant for ▶ Europa; however, cyanobacteria are
incoming radiant ▶ energy (Bhaya et al. 2000). not common components of polar seas.
In addition, many cyanobacteria are exposed to Simulation of conditions on other planets or
environmental conditions where continuous ultra- during interplanetary transport is another
violet radiation (UVR) plays an important role. approach to the estimation of limits of survival.
UVR can lead to direct photochemical damage Chroococcidiopsis sp. is able to survive in Mars-
and degradation of cellular components or to indi- like conditions when it is covered by at least
rect effects produced by reactive oxygen species. 1 mm of Mars soil analogue that reduces the
Cyanobacteria possess four lines of defense incoming ▶ UV radiation. Due to its resistance,
against UVR (Vincent 2000): they can avoid this cyanobacterium was proposed as a pioneer
UVR injuries by (1) selection of ▶ habitat (move microorganism for Mars terraforming
to deeper water, live beneath rock surface, or live (Friedmann and Ocampo-Friedmann 1995). The
deep within microbial mats); (2) production of resistance of the halotolerant Synechococcus to
umbrella compounds that filter UVR, such as the space conditions was evaluated during the
black or dark pigment Scytonemin (absorbance at BIOPAN-3 mission and the effect of space radi-
ca. 390 nm) and mycosporine-like amino acids ation on primary producers, including
(absorbance at 310–360 nm); (3) production of cyanobacteria, as part of the EXPOSE-R mission
carotenoids (canthaxanthin, myxoxanthophyll, launched on 2008 (Rabbow et al. 2009).
and related compounds), which protect the cells
from the oxidative stress caused by the UVR; and
(4) further protection against long-term effects of Future Directions
UVR exposure due to the ability to identify and
repair the photochemical damage to DNA or the As mentioned, cyanobacteria represent the earli-
photosynthetic apparatus (Vincent 2000). est organisms capable of oxygenic photosynthe-
sis. Together with other non-oxygenic
photoautotrophs, such as sulfur bacteria, they
Key Research Findings provide insight into the origin and evolution of
photosynthetic processes and data for the estima-
Some extreme environments on Earth are used as tion of their possible modifications on planets
field analogues of extraterrestrial conditions, orbiting stars of different spectral classes
allowing the limits of survival of various (Xiong and Bauer 2002).
594 Cyanobacteria
See Also Castenholz RW (1992) Species usage, concept, and evo-

lution in the cyanobacteria (bluegreen algae). J Phycol
28:737–745
▶ Adaptation Comte K, Šabacká M, Cadel S, Elster J, Komárek J (2007)
▶ Antarctica Relationship of selected Oscillatorian cyanobacteria
▶ Bacteria isolates from the Arctic and the Antarctic. FEMS
▶ Biofilm Microbiol Ecol 59(2):366–376
de Pablo MA, Pacifici A, Komatsu G (2008) A possible
▶ Carbon Cycle, Biological small frozen lake in Utopia Planitia, Mars. In: 39th
▶ Chlorophylls lunar and planetary science conference, (lunar and
▶ Chloroplast planetary science XXXIX), League City, p 1057
▶ Colonization, Biological Elster J, Benson EE (2004) Life in the polar terrestrial
environment a focus on algae and cyanobacteria. In:
▶ Cryosphere Fuller B, Lane N, Benson EE (eds) Life in the frozen
▶ Cryptoendolithic state. Taylor and Francis, London, pp 111–149
▶ Desiccation Elster J, Seckbach J, Vincent WF, Lhotský O (eds)
▶ Earth’s Atmosphere, Origin and Evolution of (2001) Algae and extreme environments; ecology and
physiology. Nova Hedwigia, Berlin, p 602
▶ Endolithic Friedmann EI (1982) Endolithic microorganisms in Ant-
▶ Endosymbiosis arctic cold desert. Science 215:1045–1053
▶ Epilithic Friedmann EI, Ocampo-Friedmann R (1995) A primitive
▶ Extreme Environment cyanobacterium as pioneer microorganism for
terraforming Mars. Adv Space Res 15:143–246
▶ Extremophiles Knoll AH (2008) Cyanobacteria and earth history. In:
▶ Fossil Herrero A, Flores E (eds) The cyanobacteria: molecu-
▶ Halophile lar biology, genomics and evolution. Caister Aca-
▶ Halotolerance demic, Norfolk, pp 1–19
Komárek J, Anagnostidis K (1998) Cyanoprokaryota
▶ Mars Analogues 1. Teil: chroococcales. In: Ettl H, Gärtner G,
▶ Membrane Heynig H, Mollenhauer D (eds) S€ usswasserflora von
▶ Mesophile mitteleuropa 19/1. Gustav Fisher, Jena, Stuttgart/
▶ Microbial Mats L€ubeck, Ulm, p 548
Komárek J, Anagnostidis K (2005) Cyanoprokaryota
▶ Microfossils 2. Teil: oscillatoriales. In: Ettl H, Gärtner G,
▶ Nitrogen Cycle, Biological Heynig H, Mollenhauer D (eds) S€ usswasserflora von
▶ Nitrogen Fixation mitteleuropa 19/2. Gustav Fisher, Jena, Stuttgart/
▶ Osmolite L€ubeck, Ulm, p 759
Komárek J, Elster J, Komárek O (2008) Diversity of
▶ pH cyanobacterial microflora of the northern part of
▶ Photosynthesis James Ross Island, NW Weddell Sea, Antarctica.
▶ Phototroph Polar Biol 31:853–865
▶ Phylogeny Oren A (2000) Salts and brines. In: Whitton BA, Potts
M (eds) The ecology of cyanobacteria; their diversity
▶ Psychrophile in time and space. Kluwer, Dordrecht/London/Boston,
▶ Stromatolites pp 281–306
▶ Thermophile Rabbow E, Horneck G, Rettberg P, Schott J-U, Panitz C,
▶ UV Radiation L’Afflitto A, von Heise-Rotenburg R, Willnecker R,
Baglioni P, Hatton J, Dettmann J, Demets R, Reitz
▶ Water Y (2009) EXPOSE, an astrobiological exposure facil-
ity on the international space station. Orig Life
Evol Biosph 39:581–598
Reed RH, Richardson DL, Warr SRC, Stewart WDP
References and Further Reading (1984) Carbohydrate accumulation and osmotic stress
in cyanobacteria. J Gen Microbiol 130:1–4
Bhaya D, Schwarz R, Grossman AR (2000) Molecular Six C, Thomas J-C, Garczarek L, Ostrowski M,
responses to environmental stress. In: Whitton BA, Dufresne A, Blot N, Scanlan DJ, Partensky F (2007)
Potts M (eds) The ecology of cyanobacteria; their Diversity and evolution of phycobilisomes in marine
diversity in time and space. Kluwer, Dordrecht/Lon- sznechococcus spp.: a comparative genomic study.
don/Boston, pp 397–442 Genome Biol 8:259
Cyanobacteria, Diversity and Evolution of 595
Strunecký O, Elster J, Komárek J (2010) Relationship in Synonyms

geographically separate Phormidium like
cyanobacteria: is there a link between north and
south polar regions? Polar Biol 33:1419–1428 Blue-green algae; Oxygenic phototrophic bacte-
Strunecký O, Elster J, Komárek J (2012a) Molecular clock ria; Oxyphotobacteria
evidence for survival of Antarctic cyanobacteria
1 (Oscillatoriales, Phormidium autumnale) from
Paleozoic times. FEMS Microbial Ecol 82:482–490
Strunecký O, Komárek J, Elster J (2012b) Biogeography Definition C
of Phormidium autumnale (Oscillatoriales,
Cyanobacteria) at western and central Svalbard. Polish Cyanobacteria are oxygenic phototrophic micro-
Polar Res 33(4):369–382 organisms. They belong to the ▶ Bacteria domain
Swingley WD, Blankenship RE, Jason R (2008) Insights
into cyanobacterial evolution from comparative geno- of life and have a plant-type ▶ photosynthetic
mics. In: Herrero A, Flores E (eds) The cyanobacteria: apparatus. They posses two photosystems, PS-I
molecular biology, genomics and evolution. Caister and PS-II, which are connected in series, and their
Academic, Norfolk, pp 21–43 reaction centers usually contain chlorophyll a.
Taton A, Grubisic S, Balthasart P, Hodgson DA,
Laybourn-Parry J, Wilmotte A (2006) Biogeographi- Water is used as the electron donor and is split
cal distribution and ecological ranges of benthic into electrons and oxygen by PS-II. The electrons
cyanobacteria in East Antarctic lakes. FEMS Microbi- are transported through an electron transport
ology Ecol 57:272–289 chain through PS-I and eventually reduce elec-
Vincent WF (2000) Cyanobacterial dominance in polar
regions. In: Whitton BA, Potts M (eds) The ecology of tron carriers. These are mainly used for the fixa-
cyanobacteria; their diversity in time and space. tion of carbon dioxide through the reductive
Kluwer, Dordrecht/London/Boston, pp 321–340 pentose phosphate pathway. Their main light-
Ward DM, Castenholz RW (2000) Cyanobacteria in geo- harvesting pigments are the blue- and
thermal habitats. In: Whitton BA, Potts M (eds) The
ecology of cyanobacteria; their diversity in time and red-colored phycobiliproteins.
space. Kluwer, Dordrecht/London/Boston, pp 37–59
Whitton BA, Potts M (2000) Introduction to the
cyanobacteria. In: Whitton BA, Potts M (eds) The Overview
ecology of cyanobacteria; their diversity in time and
space. Kluwer, Dordrecht/London/Boston, pp 1–11
Xiong J, Bauer CE (2002) Complex evolution of photo- Cyanobacteria are a monophyletic but highly
synthesis. Annu Rev Plant Biol 53:503–521 diverse group of microorganisms (Rippka
et al. 1979). Although they are often called
“blue-green algae,” they belong to the domain
Bacteria and they lack a nucleus or other cell
Cyanobacteria, Diversity and organelles (Stanier and Cohen-Bazire 1977).
Evolution of The confusion with algae, which belong to the
domain Eukarya, is understandable since
Lucas J. Stal cyanobacteria possess a plant-type photosyn-
Department of Marine Microbiology, Royal thetic system, with two photosystems, PS-I and
Netherlands Institute of Sea Research (NIOZ), PS-II, connected in series. Cyanobacteria use
Yerseke, The Netherlands water as the electron donor, which by its splitting
results in the evolution of oxygen, and their reac-
tion centers normally contain the plant pigment
Keywords chlorophyll a. Cyanobacteria fix CO2 through the
reductive pentose phosphate pathway (Calvin-
Carbon dioxide fixation; Chloroplast; Benson-Bassham cycle) with ribulose-1,5-
Cyanobacteria; Diversity; Evolution; Hetero- bisphosphate carboxylase/oxygenase (RubisCO)
cysts; Nitrogen fixation; Oxygenic photosynthe- as the CO2-fixing enzyme. This pathway is
sis; Photosynthesis also common among phototrophic Eukarya
596 Cyanobacteria, Diversity and Evolution of
(plants and algae) and other autotrophic bacteria. oxidized the massive reducing crust and the
Cyanobacteria occur in almost any illuminated euphotic layer of the ocean. Thus, it is likely
habitat on Earth, including those characterized that the origin of cyanobacteria occurred early
by extreme environmental conditions. in the evolution of life on Earth, earlier than
It is now well established that the eukaryotic 2.5 Ga.
plant cell evolved through an endosymbiotic Obviously, an oxygenic phototrophic organ-
event during which a cell engulfed a cyanobacte- ism with two photosystems did not evolve at once
rium, which subsequently evolved to become a (Olson and Blankenship 2004). There is no doubt
chloroplast, the photosynthetic factory of plant that the predecessor of the cyanobacteria was an
cells (Raven and Allen 2003). The 16S rRNA anoxygenic phototrophic organism with one pho-
genes of the chloroplast cluster phylogenetically tosystem. However, such organism might have
with the cyanobacteria. Modern cyanobacteria been morphologically indistinguishable from
enter into a large variety of symbioses with modern cyanobacteria. Some modern
microalgae and plants, often providing their cyanobacteria are capable of anoxygenic PS-I-
hosts with fixed nitrogen (Rai et al. 2000). dependent photosynthesis using sulfide as the
Cyanobacteria invented oxygenic photosyn- electron donor.
thesis and were therefore responsible for the oxy- Although monophyletic, cyanobacteria
genation of the Earth’s atmosphere (Knoll 2003). exhibit an amazing diversity. They differ two
The first big oxygen event occurred 2.4 billion orders of magnitude in size from the smallest
years ago when the oxygen concentration reached cyanobacteria measuring 0.5 mm to the biggest
10 % of its present value (GOE: Great Oxygena- of 50 mm and from unicellular forms to fila-
tion Event). Besides this rise in the oxygen level mentous species of which the trichomes can be as
of the atmosphere, there are other geological and long as 10 mm. Cyanobacteria are also quite
geochemical evidences for the presence of unique since they are among the very few organ-
cyanobacteria at that time. There are microfossils isms outside the Eukarya that display true cell
that resemble modern cyanobacteria since 2.1 Ga, differentiation. Moreover, the growth of
and there are molecular fossils such as populations of cyanobacteria may be macro-
methylhopanoids that are considered to have scopic in the form of well-defined structured
originated from them since at least 1.6 Ga, per- aggregates.
haps 2.5 Ga. Moreover, phylogenetic evidence Cyanobacteria are divided into five large divi-
also points to the presence of cyanobacteria at sions that are supported by the phylogeny of the
that time (e.g., Sanchez-Baracaldo et al. 2005). 16S rRNA gene (Fig. 1). Divisions 1 and 2 are
Recently, several geochemical studies point to unicellular. Cyanobacteria of Division 1 may
“whiffs” of oxygen before the GOE. Other stud- divide in one, two, or three planes, resulting in
ies have suggested an even earlier origin of typical morphologies of the aggregates. Division
cyanobacteria of up to almost 3.5 billion years. 2 is composed of cyanobacteria that divide by
This goes back to the earliest evidence of life on multiple fissions and produce small daughter
Earth. This evidence was based on microfossils cells, called baeocytes. Although considered uni-
interpreted to possess similarity to modern cellular, several representatives form more or less
cyanobacteria and stable isotope data suggestive regular aggregates or colonies, often surrounded
of CO2 fixation by autotrophic organisms as well with a structured sheath. Division 3 comprises all
as other organic signatures. Although these fossil filamentous cyanobacteria with solely undiffer-
traces could have been produced by other organ- entiated cells. Also these cyanobacteria may form
isms, perhaps anoxygenic photosynthesizers, various forms of more or less structured sheathed
there is no doubt that cyanobacteria must have colonies and bundles. Divisions 4 and 5 comprise
evolved long before free oxygen appeared in the filamentous cyanobacteria that show true cell dif-
atmosphere. It must have taken some time until ferentiation. Under nitrogen starvation, some
the oxygen evolved by the first cyanobacteria had cells in the filaments differentiate into
Cyanobacteria, Diversity and Evolution of 597
Cyanobacteria, Diversity and Evolution of, column: non-heterocystous filamentous cyanobacteria

Fig. 1 Diversity of Cyanobacteria and examples from (Division 3). From top to bottom: Lyngbya sp.,
each of the five divisions. Left column: heterocystous Phormidium sp., Spirulina sp., and Trichodesmium
cyanobacteria. Top three belong to Division 4: Anabaena sp. Right column: unicellular cyanobacteria. Top three
sp. with heterocysts (lighter cells) and akinetes (very large belong to Division 1: Gloeocapsa sp., the colony-forming
cells), Calothrix sp. with terminal heterocysts, and Microcystis sp., and the tiny Crocosphaera sp. Bottom:
Nodularia sp. with intercalary heterocysts. Bottom: Dermocarpa sp. with baeocytes (Division 2)
Fischerella sp. with true branching (Division 5). Middle
heterocysts (also called by the more appropriate the oxygen-sensitive nitrogenase. Many hetero-
but less common name heterocytes). Heterocysts cystous cyanobacteria also produce akinetes
have only photosystem I and therefore do not (Adams and Duggan 1999). These are differenti-
evolve oxygen and do not fix CO2. For carbon ated cells that have also been termed incorrectly
and reducing equivalents, heterocysts depend on “spores” and serve to enhance survival of the
their neighboring vegetative cells. Heterocysts organism under unfavorable conditions. Akinetes
are the site of N2 fixation. The special thick gly- are drought and radiation resistant but are not
colipid cell envelope represents a gas diffusion heat resistant like bacterial endospores. Akinetes
barrier, limiting the flux of oxygen into the cell, contain ample amounts of storage compounds,
thereby providing an anaerobic environment for such as glycogen (carbon and energy storage)
598 Cyanobacteria, Diversity and Evolution of
and cyanophycin (nitrogen storage). Akinetes vary the ratio of PEB: PUB as a response to the
germinate and form hormogonia, another form prevailing underwater light conditions (Everroad
of differentiation. Hormogonia may also differ- et al. 2006).
entiate from vegetative cells when the organism Prochloron and Prochlorothrix are
is exposed to environmental stress. Hormogonia cyanobacteria that lack phycobiliproteins and
are short, motile trichomes with cells smaller than contain chlorophylls a and b. The oceanic
those of the mature organism and do not possess picoplanktonic cyanobacterium Prochlorococcus
heterocysts. They serve as a dispersion mecha- contains the divinyl derivatives of chlorophyll a
nism for the organism and are also important for and b, which are specific for this genus (Scanlan
entering into symbiotic relationships. The main et al. 2009). Some strains contain small amounts
difference between Divisions 4 and 5 is that the of phycoerythrin, but phycobilisomes are
latter shows true branching. This occurs when a lacking, and this pigment does not serve as a
cell divides in more than one plane and distin- light-harvesting pigment in Prochlorococcus.
guishes it from apparent branching that may Another unusual cyanobacterium is represented
occur in both divisions. Heterocystous by the genus Acaryochloris which contains chlo-
cyanobacteria also may form colonies and rophyll d instead of chlorophyll a and is an adap-
aggregates. tation to the specific light conditions in its natural
Cyanobacteria are phototrophs and therefore environment.
possess photopigments. The great variety of Cyanobacteria utilize a variety of nitrogen
photopigments of cyanobacteria gives these sources (Herrero et al. 2001). Many but not all
organisms a plethora of colors and allows them cyanobacteria are capable of fixing atmospheric
to adapt to a variety of light conditions. The main dinitrogen (N2). Nitrogenase, the enzyme com-
pigment in the photosynthetic reaction centers is plex that reduces N2 to NH3, is inactivated by
the plant-type chlorophyll a. The main light- oxygen (Bergman et al. 1997). Therefore, the
harvesting pigments are the phycobiliproteins. occurrence of N2 fixation in the oxygenic
These pigments are organized in phycobilisomes cyanobacteria is paradoxical. N2-fixing
that are connected to the photosynthetic cyanobacteria have evolved mechanisms to
thylakoid membranes (Adir 2005). All ensure an anoxic intracellular environment for
phycobilisomes contain allophycocyanin. In nitrogenase. Many cyanobacteria are capable of
addition, phycocyanin (blue) or phycoerythrin fixing N2 only under anaerobic and anoxygenic
(orange to red) or both may be present, rendering conditions, a strategy that can be termed avoid-
the organism a blue-green, orange, red, or brown ance (of oxygen) (Gallon 1992). The most
color. Phycoerythrocyanin is another advanced strategy is the differentiation of hetero-
phycobiliprotein with a reddish color that has a cysts (see above). These cells contain nitroge-
rather limited distribution and occurs only in nase, and the strategy is a temporal separation
some heterocystous cyanobacteria. Phycoery- of oxygenic photosynthesis (in the vegetative
thrin comes in two forms, namely with the chro- cells) and N2 fixation (in the heterocysts). Many
mophore phycoerythrobilin (PEB) (red) and non-heterocystous N2-fixing cyanobacteria
phycourobilin (PUB) (orange). Both chromo- (filamentous as well as unicellular) exhibit a spa-
phores can be bound to phycoerythrin in various tial separation of both incompatible processes by
ratios. Some cyanobacteria are capable of com- confining the fixation of N2 to the night (Sherman
plementary chromatic adaptation (Mullineaux et al. 1998). The oceanic non-heterocystous spe-
2001). Depending on the wavelength of light, cies Trichodesmium is an enigma as it fixes N2
the amounts of phycocyanin and phycoerythrin exclusively during the daytime. It is hypothesized
will vary, turning cells blue green in red light and that this organism employs a combination of spa-
red in green light. Another form of more subtle tial and temporal separation of N2 fixation and
chromatic adaptation occurs when organisms oxygenic photosynthesis, i.e., that nitrogenase
Cyanobacterial Mats 599
activity is confined to temporary non-oxygenic Adir N (2005) Elucidation of the molecular structures of
photosynthetic cells which have also been termed components of the phycobilisome: reconstructing a
giant. Photosynth Res 85:15–32
“diazocytes.” Bergman B, Gallon JR, Rai AN, Stal LJ (1997) N2 fixation
Cyanobacteria possess a remarkably versatile, by non-heterocystous cyanobacteria. FEMS Microbiol
flexible, and reactive metabolism. In addition to Rev 19:139–185
the typical cyanobacterial metabolisms men- Everroad C, Six C, Partensky F, Thomas J-C,
Holtzendorff J, Wood AM (2006) Biochemical bases
tioned above, some cyanobacteria exhibit also of type IV chromatic adaptation in marine
C
efficient anaerobic metabolism. For instance, Synechococcus spp. J Bacteriol 188:3345–3356
they can perform anoxygenic photosystem Gallon JR (1992) Reconciling the incompatible: N2 fixa-
I-dependent photosynthesis during which sulfide tion and O2. New Phytol 122:571–609
Golden SS (2003) Timekeeping in bacteria: the
is used as the electron donor. Some species do cyanobacterial circadian clock. Curr Opin Microbiol
this in concert with oxygenic photosynthesis 6:535–540
while others rely only on anoxygenic photosyn- Herrero A, Muro-Pastor AM, Flores E (2001) Nitrogen
thesis and CO2 fixation. Cyanobacteria obviously control in cyanobacteria. J Bacteriol 183:411–425
Knoll AH (2003) The geological consequences of evolu-
possess dark metabolism in order to survive and tion. Geobiology 1:3–14
cope with the natural day and night cycle. Under Mullineaux CW (2001) How do cyanobacteria sense and
aerobic conditions, the glycogen reserve is respond to light? Mol Microbiol 41:965–971
respired through the oxidative pentose phosphate Olson JM, Blankenship RE (2004) Thinking about the
evolution of photosynthesis. Photosynth Res
cycle (the reverse of the Calvin-Benson-Bassham 80:373–386
cycle of CO2 fixation) using oxygen as the elec- Rai AN, Söderbäck E, Bergman B (2000) Cyanobacterium-
tron acceptor. Under anaerobic conditions, some plant symbioses. New Phytol 147:449–481
cyanobacteria are capable of fermenting glyco- Raven JA, Allen JF (2003) Genomics and chloroplasts
evolution: what did cyanobacteria do for plants?
gen via a variety of different pathways, and when Genome Biol 4:209
available they can use elemental sulfur as elec- Rippka R, Deruelles J, Waterbury JB, Herdman M, Stanier
tron acceptor (Stal and Moezelaar 1997). In order RY (1979) Generic assignments strain histories and
to tune all these processes to the natural day-night properties of pure cultures of cyanobacteria. J Gen
cycle, cyanobacteria are the only Bacteria with a Sánchez-Baracaldo P, Hayes PK, Blank CE (2005) Mor-
circadian clock, which occurs otherwise only phological and habitat evolution in the Cyanobacteria
within the Eukarya (Golden 2003). using a compartmentalization approach. Geobiology
3:145–165
Scanlan DJ, Ostrowski M, Mazard S, Dufresne A,
Garczarek L, Hess WR, Post AF, Hagemann M,
See Also Paulsen I, Partensky F (2009) Ecological genomics of
marine picocyanobacteria. Microbiol Mol Biol Rev
73:249–299
▶ Archean Traces of Life Sherman LA, Meunier P, Colón-López MS (1998) Diurnal
▶ Bacteria rhythms in metabolism: a day in the life of a unicellu-
▶ Belcher Group, Microfossils lar, diazotrophic cyanobacterium. Photosynth Res
58:25–42
▶ Great Oxygenation Event Stal LJ, Moezelaar R (1997) Fermentation in
▶ Microfossils cyanobacteria. FEMS Microbiol Rev 21:179–211
▶ Oxygenation of the Earth’s Atmosphere Stanier RY, Cohen-Bazire G (1977) Phototrophic pro-
▶ Photosynthesis karyotes: the cyanobacteria. Annu Rev Microbiol
31:225–274
▶ Stromatolites

Cyanobacterial Mats
Adams DG, Duggan PS (1999) Heterocyst and akinete
differentiation in cyanobacteria. New Phytol 144:3–33 ▶ Microbial Mats
600 4-Cyano-1,3-Butadiynyl
4-Cyano-1,3-Butadiynyl Cyanobutadiynyl Radical
William M. Irvine ▶ 4-Cyano-1,3-Butadiynyl

Synonyms Cyanoethane
C5N; Cyanobutadiynyl radical ▶ Ethyl Cyanide

▶ Vinyl Cyanide
Definition
The C5N ▶ radical is found in both the envelopes Cyanoethynyl Radical

of carbon stars (evolved stars whose atmospheres
contain more carbon than oxygen, the excess William M. Irvine
carbon presumably produced by helium fusion University of Massachusetts, Amherst, MA, USA
in the latter stages of the star’s life) and in cold,
dark interstellar ▶ molecular clouds (typically
those that have not been heated by star forma- Synonyms
tion). It is an intermediary in the chemistry of the
▶ cyanopolyynes and related molecules (Guelin C3N
et al. 1998). The rotational transitions of the
cyanobutadiynyl radical are observed by radio
astronomers at millimeter wavelengths. The Definition
anion of this species, C5N, has also been found
in space (Cernicharo et al. 2008). The four-atom ▶ radical C3N is found in the gas
phase in both interstellar ▶ molecular clouds and
in the expanding envelopes of evolved carbon-
See Also rich stars (evolved stars are those which have
completed their main hydrogen fusion stage and
▶ Cyanopolyyne are thus nearing the end of their energy-
▶ Molecular Cloud producing lifetimes). C3N is an intermediary in
▶ Molecules in Space the chemistry of the ▶ cyanopolyynes and related
▶ Radical radicals.
History
Cernicharo J, Guélin M, Agúndez M, McCarthy MC, The presence of C3N in the envelope of the
Thaddeus P (2008) Detection of C5N and carbon star IRC + 10216 was announced
vibrationally excited C6H in IRC +10216. Astrophys from the pattern of detected emission lines at
J 688:L83–L86
millimeter wavelengths by Guelin and Thaddeus
Guelin M, Neininger N, Cernicharo J (1998) Astronomical
detection of the cyanobutadiynyl radical C5N. Astron (1977), before the frequencies of these transitions
Astrophys 335:L1–L4 had been measured in the laboratory.
Cyanogen Radical 601
Friberg et al. (1980) stated that the chain of logic Definition

leading to this deduction was “worthy of Hercule
Poirot (Christie 1945).” This identification is a Cyanogen is a compound of formula (CN)2. It is a
particularly good example of the ability of het- colorless gas at standard temperature and pres-
erodyne (high frequency resolution) astronomi- sure. A cyanogen molecule consists of two CN
cal measurements to contribute to fundamental groups bonded together at their carbon atoms
molecular physics. (NCCN). Cyanogen is the anhydride of C
oxamide. It can be generated from cyanide com-
pounds and solutions of metal salts (such as cop-
See Also per(II) sulfate).
▶ Cyanopolyyne 2CuSO4 þ 4KCN ! ðCNÞ2 þ 2CuCN

▶ Molecular Cloud þ 2K2 SO4 (1)
▶ Radical It is also formed when ▶ nitrogen and ▶ acet-
▶ Stellar Evolution ylene are acted upon by an electrical discharge.
It is implicated as a possible phosphorylating
agent for nucleosides and a possible precursor to
References and Further Reading cyanates and ureas.
Christie A (1945) Sparkling cyanide. Collins, London

Friberg P, Hjalmarson Å, Irvine WM (1980) Interstellar
C3N: detection in taurus dark clouds. Astrophys J 241:
L99–L103
See Also
Guelin M, Thaddeus P (1977) Tentative detection of the
C3N radical. Astrophys J 212:L81–L85 ▶ Electric Discharge
Cyanogen
Henderson James (Jim) Cleaves II Cyanogen nitride

Earth-Life Science Institute (ELSI),
Tokyo Institute of Technology, Meguro-ku, ▶ Cyanamide
Tokyo, Japan
Institute for Advanced Study, Princeton,
NJ, USA
Washington, DC, USA Cyanogen Radical
of Technology, Atlanta, GA, USA William M. Irvine
Synonyms
Carbon nitride; Dicyan; Dicyanogen; Nitriloace- Synonyms

tonitrile; Oxalic acid dinitrile; Oxalonitrile;
Oxalyl cyanide CN; Cyano radical
602 Cyanogenamide
Definition Jefferts KB, Penzias AA, Wilson RW (1970) Observation

of the CN radical in the Orion nebula and W51.
Astrophys J 161:L87
The diatomic radical CN, containing carbon and McKellar A (1940) Evidence for the molecular origin of
nitrogen, is widely observed in the ▶ interstellar some hitherto unidentified interstellar lines. Publ
medium of the ▶ Milky Way and external galax- Astron Soc Pac 52:187–192
ies. It plays an important role in interstellar chem-
istry, being an intermediary in the production and
destruction of such important species as HCN and
HNC. The CN radical is also prominent in the
visible wavelength spectra of cometary comae Cyanogenamide
(atmospheres), where it is presumably a photo-
dissociation product of molecules such as HCN ▶ Cyanamide
that are sublimated from the icy nucleus. Note
that the molecule C2N2 is also referred to as the
cyanogen radical. A chemical compound that
contains the CN functional group (carbon triple
bonded with nitrogen) is called a cyanide, while
an organic compound with this CN group is Cyanomethane
called a nitrile.
▶ Acetonitrile
History
Unlike most interstellar molecules, which were

discovered by radio astronomical observations, Cyanomethanimine
CN was first identified at ultraviolet wavelengths
in 1940, and it was one of the first known inter- William M. Irvine
stellar molecular species. The fundamental rota- University of Massachusetts, Amherst,
tional line was observed radio astronomically in MA, USA
1970 by Jefferts et al.
See Also Synonyms
▶ Comet HCN dimer; HNCHCN

▶ Hydrogen Isocyanide
▶ Interstellar Medium Definition
Cyanomethanimine is an organic molecule that is
a less saturated relative of ▶ aminoacetonitrile
(glycine nitrile, NH2CH2CN). It has three ▶ iso-
References and Further Reading mers, N-, E-, and Z-cyanomethanimine.
Crovisier J, Encrenaz T (2000) Comet science: the study
Cyanomethanimine has been proposed as an
of remnants from the birth of the solar system. Cam- intermediary in the prebiotic synthesis of the
bridge University Press, Cambridge purine nucleobase adenine.
Cyclopropenylidene 603
History polyynes are organic compounds with alternating

single and triple bonds, the simplest being
E-Cyanomethanimine has recently been detected ▶ diacetylene (HCC─CCH), cyanopolyynes
by radio astronomers in the interstellar medium are end-capped by the cyano group (─CN).
(Zaleski et al. 2013). Examples are HCC─CC─CN and
HCC─CC─CC─CN.
C
See Also
See Also
▶ Adenine
▶ Aminoacetonitrile ▶ Circumstellar Chemistry
▶ Glycine ▶ Diacetylene
▶ Molecules in Space ▶ Molecules in Space
Zaleski DP, Seifert NA, Steber AL, Muckle MT, Loomis

RA, Corby JF, Martinez O, Crabtree KN, Jewell PR, Cyclic Replicator Equation
Hollis JM, Lovas FJ, Vasquez D, Nyiramahirwe J,
Sciortino N, Johnson K, McCarthy MC, Remijan AJ, ▶ Hypercycle
Pate BH (2013) Detection of E-cyanomethanimine
toward Sagittarius B2(N) in the green bank telescope
PRIMOS survey. Astrophys J 765:L10–L15
Cyanomethylamine Cyclohexa-1,3,5-triene
▶ Aminoacetonitrile ▶ Benzene
Cyanophyceae
Cyclopropenylidene
▶ Cyanobacteria
William M. Irvine
Cyanopolyyne
Steven B. Charnley Synonyms

Solar System Exploration Division, Code 691,
Astrochemistry Laboratory, NASA Goddard C3H2, c-C3H2
Definition
Definition
The 3-carbon ring molecule C3H2 is classified
Cyanopolyynes are long carbon-chain molecules chemically as a ▶ carbene and is highly reactive
found in many astrochemical sources. Whereas in the laboratory. It was the first cyclic molecular
604 Cysteine
species detected in interstellar ▶ molecular Cernicharo J, Cox P, Fosse’ D, Gusten R (1999) Detection
clouds. Immediately following its identification of linear C3H2 in absorption toward continuum
sources. Astron Astrophys 351:341–346
at short radio wavelengths, it was discovered to Matthews HE, Irvine WM (1985) The hydrocarbon ring
be nearly ubiquitous in the ▶ interstellar medium C3H2 is ubiquitous in the galaxy. Astrophys J 298:
(Matthews and Irvine 1985), and it has subse- L61–L64
quently also been found in the envelope of Thaddeus P, Vrtilek JM, Gottlieb CA (1985) Laboratory
and astronomical identification of cyclopropenylidene,
evolved carbon-rich stars (evolved stars are C3H2. Astrophys J 299:L63–L66
those which have completed their main hydrogen
fusion stage and are thus nearing the end of their
energy-producing lifetimes). The linear ▶ isomer
of C3H2 is also detected in molecular clouds, but
its abundance is typically an order of magnitude Cysteine
lower than that of the cyclic isomer (Cernicharo
et al. 1991). In diffuse clouds, the abundance ratio Kensei Kobayashi
between the linear and the cyclic isomers Yokohama National University, Tokiwadai,
increases by a factor 10 with respect to that Hodogayaku, Yokohama, Japan
observed in dense molecular clouds (Cernicharo
et al. 1999). Both 13-carbon and deuterated iso-
topic forms of C3H2 have been detected
astronomically. Definition
Cysteine is one of the 20 protein ▶ amino acids,

History whose structure is shown in Fig. 1. Its three-
letter symbol and one-letter symbol are Cys
The laboratory measurement of the frequencies and C, respectively. Among the protein amino
of several C3H2 rotational transitions by acids, only cysteine and methionine contain a
Thaddeus et al. (1985) proved the identification sulfur atom. The side chain of cysteine contains
of emission lines that they had earlier observed a thiol (-SH) group, which is often present in
astronomically. the active site of enzymes. In proteins, two
cysteine residues often form a disulfide bond
(▶ Cystine) which is essential in stabilizing the
See Also protein 3D structure. Cysteine has been pro-
duced in a variety of prebiotic experiments
▶ Carbene from reducing gas mixtures but has not been
▶ Deuterium detected in carbonaceous chondrites, possibly
▶ Isomer due to its instability.
▶ Molecular Cloud
References and Further Reading Cysteine, COOH

Fig. 1 Chemical structure
of cysteine H2N C H
Cernicharo J, Gottlieb CA, Guelin M, Killian TC,
Paubert G, Thaddeus P, Vrtilek JM (1991) Astronomi- CH2
cal detection of linear H2CCC. Astrophys J 368:
L39–L42 SH
Cytochromes 605
See Also See Also
▶ Amino Acid ▶ Amino Acid

▶ Cystine ▶ Cysteine
▶ Proteins, Primary Structure ▶ Disulfide Bond
▶ Protein
▶ Thiol C
Cystine Cytochromes
Kensei Kobayashi Juli Peretó

Yokohama National University, Tokiwadai, Institut Cavanilles de Biodiversitat i Biologia
Hodogayaku, Yokohama, Japan Evolutiva, Universitat de València, València,
Spain
Cystine (Fig. 1) is formed by the oxidative con- Cytochromes are a class of electron-transferring
densation of two ▶ cysteine molecules. Cysteine, metalloproteins containing a heme as prosthetic
one of the 20 ▶ protein amino acids, has a ▶ thiol group and that participate in many different respi-
(─SH) group. In aqueous solution, two thiol ratory and photosynthetic ▶ electron transport
groups readily oxidize to form a ▶ disulfide chains, usually as membrane-bound electron car-
bond (─S─S─). Thus, cystine rather than cyste- riers. The function of cytochromes as electron
ine is usually determined when the amino acid carriers involves the alternate ▶ oxidation and
composition of protein hydrolysates is analyzed. ▶ reduction of the iron ion present in the heme
It can be easily reduced to give two cysteine group, one electron each time (i.e., between the
molecules. In protein molecules, cysteine resi- reduced ferrous state and the oxidized ferric
dues make intramolecular disulfide bonds, state), and with a standard ▶ redox potential
which stabilize protein tertiary structure. between 100 and +500 mV. The cytochromes
Cystine, Fig. 1 Chemical O

structure of cystine
NH2 S OH
HO S NH2
O
606 Cytoplasm
are classified on the basis of their characteristic,

redox-sensitive, visible absorbance spectra. Cytosine

See Also Earth–Life Science Institute (ELSI), Tokyo
Institute of Technology, Meguro–ku, Tokyo, Japan
▶ Electron Carrier Institute for Advanced Study, Princeton, NJ, USA
▶ Electron Transport Blue Marble Space Institute of Science,
▶ Oxidation Washington, DC, USA
▶ Photosynthesis Center for Chemical Evolution, Georgia Institute
▶ Redox Potential of Technology, Atlanta, GA, USA
▶ Reduction
▶ Respiration
Definition
Cytosine (C) is one of the four heterocyclic nitrog-

enous bases found in DNA (A, T, C, and G) and
Cytoplasm
RNA (A, U, C, and G). It is a pyrimidine with two
functional group substituents: an amine at the C4
Juli Peretó
position and a keto group at the C2 position. When
cytosine is combined with ribose via a glycosidic
linkage between its N1 nitrogen and the C1 position
Spain
of the sugar, it forms a nucleoside called cytidine;
removal of the 20 OH group of this molecule results
in the formation of 20 -deoxycytidine also known as
Definition
deoxycytidine. In Watson-Crick base pairing in
nucleic acids, these derivatives form three hydro-
Cytoplasm is the internal space of a ▶ cell
gen bonds with guanine.
containing all soluble chemical components
Cytosine and its derivatives hydrolyze fairly
and, in the case of eukaryotic cells, the organelles
rapidly under physiological conditions to give ura-
(▶ nucleus, mitochondria, plastids, microbodies,
cil via deamination, with a half-life of approxi-
etc.), the cytoskeleton, and the endomembrane
mately 73 years at 37 C at pH 7. In biological
system (endoplasmic reticulum, Golgi apparatus,
systems, this relatively rapid loss of structural
etc.). The fluid, non-particulate fraction of the
genetic information is corrected by DNA repair
cytoplasm is known as cytosol.
enzymes.
Cytosine has been synthesized under simulated
prebiotic conditions from cyanoacetaldehyde
See Also
and urea as well as from ▶ cyanoacetylene and
cyanate. It can also be derived from the deamina-
▶ Cell
tion of 2,4-diaminopyrimidine, itself derived
▶ Cell Membrane
from the condensation of guanidine with
▶ Nucleus
cyanoacetaldehyde.
See Also
Cytoplasmic Membrane
▶ Cyanoacetylene
▶ Cell Membrane ▶ Pyrimidine Base
▶ Membrane ▶ Uracil (Ura)
D
D/H fraction of a d-enantiomer over the corresponding

L-enantiomer. Abiotically synthesized chiral
▶ Deuterium/Hydrogen Ratio molecules have a D/L-ratio of 1, or, in other
words, they occur as a racemic mixture, as long
as no enantiomerically enriched reactants
or catalysts are used. In biology, however,
D/L-Ratio amino acids and sugars show a D/L-ratio that is
unequal to 1. The D/L-ratio of amino acids in
Uwe J. Meierhenrich ▶ proteins is nearly zero, whereas the D/L-ratio
Institut de Chimie de Nice (ICN), University of ribofuranosyl sugar molecules in ▶ DNA is
Nice-Sophia Antipolis, Nice, France infinite.
Keywords Overview
Amino acids; Biomolecular asymmetry; Chiral- A molecular structure that is non-superimposable

ity; Homochirality; Sugars with its mirror image is said to be chiral (see
▶ chirality). Similar to the right and the left
hand, chiral molecules exhibit identical physico-
Synonyms chemical properties such as molecular weight,
size, and color. Corresponding chiral molecules
Enantiomeric ratio are called enantiomers and their resolution in the
laboratory is often challenging. In particular for
chiral amino acids and chiral sugars, the D,L
Definition nomenclature is still in use, which distinguishes
between right-handed D-amino acids and left-
Chiral molecules such as amino acids and sugars handed L-amino acids or right-handed D-sugars
are composed of enantiomers that occur in and left-handed L-sugars. The synthesis of amino
stereochemical D- or L-configurations or in a mix- acids in the laboratory, via the Strecker mecha-
ture of them. The D/L-ratio determines the nism or via the Miller-Urey experiment, leads to a

DOI 10.1007/978-3-662-44185-5
608 D-Amino Acids
racemic mixture of the amino acids with a D/L-

ratio of exactly 1. Also, the formose reaction D-Amino Acids
produces sugar molecules in a racemic ratio,
that is, the D/L-ratio equals 1. However, living Henderson James (Jim) Cleaves II
organisms tend to prefer one conformation of Earth–Life Science Institute (ELSI), Tokyo
the two possible enantiomers. Proteins are exclu- Institute of Technology, Meguro–ku, Tokyo,
sively composed of L-amino acids. The sugars Japan
occurring in RNA and DNA are exclusively of Institute for Advanced Study, Princeton, NJ,
the D-configuration. The genetic material of bio- USA
logical organisms does not use their mirror-image Blue Marble Space Institute of Science,
L-sugar structures; thus, the molecular symmetry Washington, DC, USA
in living organisms is broken. Today it remains Center for Chemical Evolution, Georgia Institute
unknown how the biomolecular asymmetry was of Technology, Atlanta, GA, USA
triggered. Various hypothetical models exist to
explain biomolecular asymmetry, among them
(a) asymmetric photochemistry and the interac- Definition
tion of racemic organic molecules with circularly
polarized light, (b) asymmetric phase transitions D-amino acids are a-amino acids that have the
involving the precipitation of enantioenriched or R configuration at their a-carbon atom, with
enantiopure crystals or asymmetric adsorption the exception of cysteine, which is of the
processes, or (c) the weak nuclear interaction L configuration. D-amino acids are produced by
and the transfer of its inherent asymmetry to enzymatic post-translational modification of the iso-
chiral organic molecules. The discussion of the meric L-amino acids which are ribosomally encoded.
validity of these hypotheses is influenced by the D-amino acids are found in some bacterial cell walls
detection of D/L-ratios between 0.8 and 1 for dif- as well as in the some fungal antibiotic peptides.
ferent amino acids in various meteorites. Until
now, L-enantiomeric excesses of up to 18.5 %
See Also
have been identified in the Murchison meteorite
but an ▶ amino acid with a D/L-ratio larger than
▶ Amino Acid
1 has never been detected.
See Also
Dark Cloud
▶ Amino Acid
▶ Asymmetric Reaction, Absolute ▶ Dust Cloud, Interstellar
▶ Carbohydrate
▶ Carbonaceous Chondrites, Organic Chemistry of
▶ DNA
▶ Homochirality Dark Fringe Interferometry
▶ Polarized Electron
▶ Polarized Light and Homochirality ▶ Nulling Interferometry
▶ Protein

Dark Reactions
Meierhenrich U (2008) Amino acids and the asymmetry of
life. Springer, Heidelberg ▶ Calvin-Benson Cycle
Darwin’s Conception of the Origins of Life 609
History
Dark Streaks (Mars)
Charles Darwin (1807–1882) is the author of the
Alessandro Airo principal evolutionary theory of the nineteenth
Institut f€
ur Geologische Wissenschaften century, which founded the most important part
Tektonik und Sedimentäre Geologie, Freie of the evolutionary thought of the twentieth
Universität Berlin, Fachbereich century.
Geowissenschaften, Berlin, Germany The son of a physician, he abandoned his own
medical studies in Edinburgh after 2 years and D
went to Cambridge to study theology. There, he
Definition was initiated into botany and geology. In 1831, he
embarked on the Beagle for a 5-year trip around
Dark streaks is a general term for temporarily the world. As the only geologist on board, he had
darkened surface patches occurring on to study the formation of the atoll island; more-
various slopes on Mars (e.g., crater walls or over, he did the broad work of a naturalist, study-
dune faces) that display a range of morphologies ing also animals and plants. He came back to
(e.g., conical, linear, or braided). The darkening England as an acknowledged geologist. How-
of the surface is believed to be caused by ever, he and his family settled in Down, in the
the removal of the surficial dust cover due to southwest of London, where he prepared, for
dry grain flow, wetting of the soil, or fluvial 20 years, a theory about the origin of species. In
activity. Over the course of months and 1858, when he received a letter from Wallace, a
years, the dark coloration fades away, which is naturalist living in Malaysia, he decided to pub-
presumably the result of the surficial accumula- lish his book rapidly. On the Origin of Species by
tion of bright dust particles and/or drying of Means of Natural Selection, or the Preservation
the soil. of Favoured Races in the Struggle for Life was
published in 1859, and it expounded his theory
based on the variation and selection of living
beings.
See Also
In the first issue of his book, Darwin claimed
that the tree of life came from a unique and
▶ Slope Lineae, Recurrent
simple ancestor. However, the lack of fossils
▶ Slope Streaks (Mars)
from the oldest period of Earth led Darwin to be
very careful on this topic. In the first edition of his
book, he only wrote a few words about the origin
of life in the last lines of the last chapter:
Darwin’s Conception of the Origins
There is a grandeur in this view of life, with its
of Life several powers, having been originally breathed
into a few forms or into one; and that, whilst this
Stéphane Tirard planet has gone cycling on according to the fixed
Centre François Viète d’Histoire des Sciences law of gravity, from so simple a beginning endless
forms most beautiful and most wonderful have
et des Techniques EA 1161, Faculté des been, and are being, evolved.
France Doing some concessions on the social context,
he changed these lines in the second edition, but
he continued to assume that there was a very
Keywords simple common ancestor to all the living beings.
However, Darwin never published any text
Evolution; Natural selection; Variation about the process of the formation of the first
610 Darwinian Approaches
living beings. He explained his position in a letter

to his friend, the botanist Joseph Dalton Hooker Daughter Molecule, Comet
(1817–1911):
It is often said that all the conditions for the first Therese Encrenaz
production of a living organism are now present, LESIA, Observatoire de Paris - Section de
which could have been present. But if (and oh what Meudon, Meudon, France
a big if) we could conceive in some warm little
pond with all sort of ammonia and phosphoric
salts, – light, heat, electricity &c. present, that a
protein compound was chemically formed, ready Definition
to undergo still more complex changes, at the pre-
sent day such matter would be instantly devoured, A daughter molecule is a molecule produced
or absorbed, which would not have been the case
before living creatures were formed. (Calvin 1969) from the photodissociation of a parent molecule
which is directly outgassed from a cometary
In this quote, Darwin underlined several nucleus. The most abundant parent molecule is
important points directly related to his evolution H2O; its dissociation products are H and
theory. Firstly, he claimed the possibility of a OH. Other parent molecules include CO, CO2,
complexification of matter before the existence CH4, NH3, HCN, H2CO, and H2S. Daughter mol-
of life, and he indicated possible chemical ways. ecules include CN, CS, and CO (which appears to
Secondly, he analyzed the particular case of the be both a parent and a daughter molecule).
origin of life and suggested that life itself Daughter molecules and radicals are mostly
prevented a new formation of life. Therefore, detected from their visible and ultraviolet spectra,
according to Darwin, the origin of life was the while parent molecules are better observed with
unique event in the history of life that could not IR and millimeter spectroscopy.
repeat itself.
See Also
See Also ▶ Comet

▶ Parent Molecule, Comet
▶ Huxley’s Conception on Origins of Life
▶ Lamarck’s Conception of Origins of Life
De Duve, Christian
Stéphane Tirard
Calvin M (1969) Chemical evolution. Molecular evolu- Centre François Viète d’Histoire des Sciences
tion towards the origin of living systems on earth and et des Techniques EA 1161, Faculté des
elsewhere. Clarendon Press, Oxford
Darwin C (1852) The origins of species. Penguin Books,
London, 1985 France
Christian de Duve was a Belgian medical doctor

and biologist. In 1974, he was awarded with the
Darwinian Approaches Nobel Prize of medicine and physiology for his
work and discoveries of lysosomes and
▶ Genetic Algorithms peroxisomes.
Deamination 611
Throughout his lifelong career, he develops a phenomenon. Life first developed itself in seas
rich philosophical thought notably regarding the and then on continents. Therefore, the Indian
origin of life, publishing several number of books man, in fact de Maillet, argues against the reli-
on that subject. He claimed that chemistry deals gious discourse.
with deterministic phenomena and consequently
that the origin of life depended on the initial
conditions in which it arose. Therefore, he See Also
claimed that there are probably numerous cases
of life in the universe. ▶ Buffon’s Conception of Origins of Life D
Moreover, he distinguished two sorts of evo- ▶ Spontaneous Generation, History of
lution: the horizontal evolution that is responsible
of diversity of species and depends of environ-
ment and the vertical evolution that is responsible
for the increase of complexity of organisms and
Dead Zone
does not depend of environment. For example,
for him, the development of the brain is indepen-
▶ Oxygen-Minimum Zone
dent of the environmental conditions. For that
reason, de Duve claimed that the emergence of
the human species was not so improbable.
Deamination
De Maillet’s Conception of Origins Matthew Levy

of Life Michael F. Price Center, Albert Einstein College
of Medicine, Bronx, NY, USA
Stéphane Tirard
et des Techniques EA 1161, Faculté des Keywords
France Ammonia; Degradation; Hydrolysis
History Definition
At the end of the seventeenth century, Benoı̂t de Deamination describes the loss of an amine group
Maillet (1656–1738) wrote a book entitled from a molecule.
Telliamed, ou entretiens d’un philosophe indien
avec un missionnaire français (La Haye chez
Pierre Gosse) and signed under this pseudonym: Overview
Telliamed. This book was published in 1748 and
1755 after de Maillet’s death. It presents an Deamination is likely to have played an impor-
imaginary “entretien” between a missionary and tant role in the availability of the biological build-
an Indian man, who explains his conception of ing blocks of life. In the case of the nucleobases,
the world and of life. With this astucious rhetor- the components of ▶ RNA and ▶ DNA, the dom-
ical form, de Maillet presented his own view inant decomposition products of their exocyclic
through the Indian man’s mouth. According to amine-bearing bases ▶ adenine, ▶ guanine, and
him, life on earth came from a panspermic ▶ cytosine are the deaminated products
612 Debris Disk
hypoxanthine, xanthine, and uracil, respectively. References and Further Reading

A calculation of the half-lives for the rates of
decomposition of these bases shows that these Bada JL, Miller SL (1968) Ammonium ion concentration
in the primitive ocean. Science 159(813):423–425
compounds are not stable on a geologic timescale
Frederico LA, Kunkel TA, Shaw BR (1990) A sensitive
at temperatures much above 0 C. This has led to genetic assay for the detection of cytosine deamina-
the suggestion that life would be more likely to tion: determination of rate constants and the activation
evolve under cold conditions, where biologically energy. Biochemistry 29(10):2532–2537
Levy M, Miller SL (1998) The stability of the RNA bases:
important material can accumulate, rather than at
implications for the origin of life. Proc Natl Acad Sci
elevated temperatures such as hydrothermal U S A 95(14):7933–7938
vents, where they would be rapidly destroyed Lindahl T, Nyberg B (1974) Heat-induced deamination of
(Levy and Miller 1998). cytosine residues in deoxyribonucleic acid. Biochem-
istry 13(16):3405–3410
Deamination is important for other biomole-
cules such as amino acids. In particular, the
deamination of aspartic acid to fumaric acid and
▶ ammonia has been shown to have a half-life of Debris Disk
2.8 107 years at 0 C and 96,000 years at 25 C
(Bada and Miller 1968). Equilibrium measure- Paul Kalas
ments for this deamination reaction have previ- Astronomy Department, University of California,
ously been utilized to estimate the concentration Berkeley, CA, USA
of ammonium ions on the early Earth (Bada and
Miller 1968). Thus, deamination is not just a
route to the destruction of biomolecules, but Keywords
also a potential tool to ascertain conditions in
the oceans of the early Earth or other Extrasolar planetary systems; Interplanetary dust
environments. grains
In biology, one of the most critical deamina-
tion reactions is the hydrolysis of cytosine to
uracil and ammonia. At neutral pH (7) and Synonyms
37 C, the half-life of cytosine through deamina-
tion is 200 years in single-stranded DNA and Planetary debris disks; Protoplanetary disk of
30,000 year in double-stranded DNA (Lindahl second generation
and Nyberg 1974; Frederico et al. 1990). Mea-
surements for the rate of deamination of free
cytosine are similar, albeit slightly faster than Definition
those reported for single-stranded DNA
(60 years at 37 C; (Levy and Miller 1998)). A debris disk is a flattened distribution of dust
This deamination rate is sufficiently fast that bio- surrounding a main sequence star that is continu-
logical systems must have evolved repair mech- ously replenished by the collisional erosion of
anisms fairly early in their history to prevent the asteroids and comets. A debris disk may be pre-
resulting DNA damage. sent for much of the lifetime of a star, as is
evident from the presence of the Solar System’s
zodiacal cloud (▶ Zodiacal Light).
See Also
History
▶ Amino Acid
▶ Ammonia Interplanetary dust particles were first observed
▶ Nucleic Acids as the naked eye phenomenon of ▶ zodiacal light
Debris Disk 613
Debris Disk, Fig. 1 The debris belt surrounding the System. Light from the first magnitude star has been
bright, nearby A3V star Fomalhaut is detected in optical blocked by a coronagraph. The planet ▶ Fomalhaut b is
scattered light using the Hubble Space Telescope (Kalas detected just 18 AU inside of the belt’s inner edge, and
et al. 2008). For scale, the yellow ellipse indicates a 30 AU counterclockwise orbital motion is detected from obser-
orbit that would be analogous to Neptune in our Solar vations at two epochs (2004 and 2006)
in our own solar system. In 1983, the ▶ Infrared Mauna Kea, Hawaii. These data showed that cir-
Astronomical Satellite (IRAS) began an all-sky cumstellar dust disks showed various
imaging survey at 12, 25, 60, and 100 mm. asymmetries along radial and azimuthal direc-
Approximately 15 % of bright main sequence tions (Holland et al. 1998; Greaves et al. 1998).
stars revealed excess thermal emission, typically Beta Pictoris was also shown to be asymmetric in
at 60 mm, that was attributed to thermal emission the radial and vertical directions (Lagage and
from circumstellar dust grains. Stellar Pantin 1994; Kalas and Jewitt 1995; Burrows
coronagraphy and imaging showed a relatively et al. 1995). These results were the basis of the-
bright and extended dust disk surrounding the oretical investigations on how planetary compan-
nearby A star Beta Pictoris (Smith and Terrile ions could govern the structure of circumstellar
1984). An excess of infrared emission due to dust disks (Roques et al. 1994; Ozernoy et al.
circumstellar dust was also called the “Vega phe- 2000). Planetary companions were subsequently
nomenon,” since the first IRAS discovery was detected around Fomalhaut (Kalas et al. 2008;
first made for the bright star Vega (Aumann Fig. 1), HR 8799 (Marois et al. 2008) and Beta
et al. 1984). The Vega phenomenon was attrib- Pictoris (Lagrange et al. 2009).
uted to the same dust replenishment mechanisms
that create the zodiacal light – the collisional
erosion of comets and asteroids that formed Overview
around other stars. The essential timescale argu-
ment was that collisions, radiation pressure, and The study of debris disks aims to characterize the
▶ Poynting-Robertson drag limit the lifetime of evolution of planetary systems with respect to the
circumstellar dust grains to less than 105 years, population of minor bodies such as comets and
whereas the ages of the Vega phenomenon stars asteroids. Debris disks are distinct from primordial
were all greater than ten million years. The IRAS circumstellar disks that are observed in star-
mission was succeeded by the launches of the forming regions such as Orion and Taurus
▶ Infrared Space Observatory (ISO, 1995) and Aurigae. The term “debris disk” generally refers
the Spitzer (2003) and Herschel (2009) space to dust observed at mid-infrared wavelengths
telescopes. Spatially resolved imaging was most around stars old enough that their gaseous proto-
successful at 350–850 mm wavelength from the planetary disks have mostly or completely dissi-
ground, particularly with the SCUBA instrument pated. Debris disks are typically greater than
on the 15-m James Clerk Maxwell Telescope on 10 Myrs old and contain significantly less gas
614 Debris Disk
and total mass than primordial disks, and planets Thermal (far)-infrared observations offer relatively
within a debris disk have finished forming and are good estimates of grain temperature, radial loca-
in the process of evolving their atmospheres and tion, and total mass, with some inferences for grain
surfaces. An external observer would also find that size and composition. In contrast, scattered light
our planetary system has a debris disk in the inner observations give the radial, azimuthal, and vertical
Solar System (zodiacal light and asteroid belt) and distribution of grains, as well as estimates of grain
the outer solar system (see ▶ Kuiper Belt), size, composition, and porosity by analyzing wave-
corresponding to the locations where the colli- length dependence and polarization.
sional erosion of comets and asteroids replenishes The fundamental parameter in debris disk sci-
interplanetary dust particles. Since the cumulative ence is the ratio of the infrared luminosity of the
surface area of a planetary system is dominated by dust relative to the total, bolometric luminosity of
dust grains (the total mass is dominated by the the host star (Ldust/Lstar). Since debris disks are
largest planets), debris disks are the easiest com- optically thin, the fractional infrared luminosity
ponent of a planetary system to directly detect in is a measure of total dust mass. For scattered
reflected light or thermal emission. One of the light images, the measured surface brightness as
most important qualities of debris disks is that a function of distance from the star gives the radial
the gravitational influence of planets disturbs the decrease of grain number density. Other parame-
overall structure of a disk. Therefore debris disks ters often derived in the literature are the inner and
can flag a system for the presence of planets long outer radii of a debris disk, the grain temperature,
before the planets can be detected by other the grain size distribution, and the grain size below
methods (Kuchner and Holman 2003). which grains will be removed from the system by
radiation pressure (the grain blowout size). For
example, around luminous A stars such as Beta
Basic Methodology Pictoris and Fomalhaut, grains smaller than
5–10 mm are ejected by radiation pressure.
The successful detection of solid objects next to Spectroscopy has been used to infer not only
bright stars is not only a question of angular resolu- the composition of constituent grains but also
tion but also one of contrast. Debris disks were first their dynamics. Overall, spectra of debris disks
detected using thermal infrared techniques (>3 mm) reveal grain properties very similar to solar sys-
because the contrast between the bright star and the tem comets, such as crystalline silicate and oliv-
faint disk is less in the infrared than at optical ine emission features, as well as atomic gas such
wavelengths: whereas a star has an energy spectrum as neutral Fe and Ca+.
that peaks in the optical and declines in the infrared, Another key component of debris disk science
the energy spectrum of dust grains peaks at infrared is estimating the ages of the host stars in order to
wavelengths, due to the much cooler temperature of understand the evolution of planetary systems.
the dust. Therefore infrared and submillimeter Stellar age indicators such as X-ray emission and
detectors are most capable of separating the radia- the presence of lithium (destroyed as stars age) are
tion emitted from the star from the radiation emitted particularly robust for the low-mass (late-type)
by dust grains. The disadvantage is that since resolv- stars. However, since main sequence stars have
ing power scales as wavelength over the telescope long departed from the cluster environment in
diameter (D), the longer the wavelength used, the which they formed, the method for physically
larger the telescope required. associating the low-mass stars with the high-
At optical and near-infrared wavelengths, debris mass stars is by analyzing galactic space motions,
disks are detected by grains reflecting (scattering) thereby identifying groups of stars with common
light from the star, rather than by the dust’s own velocities, particularly in the azimuthal direction
emission. Star-disk contrast is a greater challenge around the galaxy. Data from the ▶ Hipparcos
than at far-infrared wavelengths, but the angular mission has been central in the identification of
resolution achievable is significantly greater. stellar “moving groups” or “kinematic groups.”
Debris Disk 615
Key Research Findings et al. 2009). These results give further evidence
that debris disk radial structure is governed by the
Approximately 15 5 % of stars have debris dynamics of planets, with direct analogies to our
disks, as suggested by the infrared surveys own Solar System.
conducted with IRAS, ISO, and the ▶ Spitzer
Space Telescope. The stellar age estimates indi-
cate that stars with debris disks happen to be the See Also
younger (10–100 Myr) stars in the ▶ solar neigh-
borhood. For example, the Beta Pictoris moving D
▶ Coronagraphy
group has several dozen candidate members ▶ Dust Grain
(Song et al. 2003), and among these there are ▶ Fomalhaut b
four debris disks that have been imaged in ▶ HR 8799: The First Directly Imaged Multi-
scattered light, though with significantly different planet System
properties such as mass, suggesting that disk evo- ▶ Infrared Excess
lution is a stochastic process (Kalas et al. 2007). ▶ Interplanetary Dust Particle
Several mechanisms have been proposed to ▶ Late Heavy Bombardment
explain the large variations in dust mass, such ▶ Planetesimals
as stochastic collisions between large ▶ planetes- ▶ Protoplanetary Disk
imals and dynamical stirring as planets grow and ▶ Protoplanetary Disk of Second Generation
dynamically evolve (Wyatt 2008). Despite a sig- ▶ Snow Line
nificant variation in debris disk properties at a ▶ Zodiacal Light
given stellar age, the general finding is that debris
disk dust mass varies approximately inversely
with age or with age squared (Zuckerman 2001; References and Further Reading
Rieke et al. 2005).
Aumann HH, Beichman CA, Gillett FC et al (1984) Dis-
Most debris disks have inner clearings of covery of a shell around Alpha Lyrae. Astrophys J 278:
material approximately 10–100 AU in radius L23–L27
(Zuckerman and Song 2004; Kalas et al. 2006). Backman D et al (2009) Epsilon Eridani’s planetary debris
In a few cases this corresponds to the “ice line,” disk: structure and dynamics based on Spitzer and
CSO observations. Astrophys J 690:1522–1538
but in many other cases the most likely explana- Burrows CK, Krist JE, Stapelfeldt KR et al (1995) HST
tion is dynamical clearing by a planetary system. observations of the beta Pictoris circumstellar disk.
For example, the debris disk surrounding Bull Am Astron Soc 187:3205
Fomalhaut is a torus of dust beginning 133 AU Greaves JS et al (1998) A dust ring around epsilon Eridani:
analog to the young solar system. Astrophys J 506:
from the star with a Jupiter mass planet imaged L133–L137
just 18 AU interior to the belt (Fig. 1; Kalas Hillenbrand et al (2008) The complete census of 70-micron
et al. 2008). bright debris disks. Astrophys J 677:630–656
In many cases, the far-infrared spectral energy Holland WS et al (1998) Submillimetre images of dusty
debris disks around nearby stars. Nature 392:788–791
distributions cannot be fit by single components Kalas P, Jewitt D (1995) Asymmetries in the beta Pictoris
of dust, but often require at least one warm inner dust disk. Astron J 110:794–804
component and one cold outer component with Kalas P et al (2006) First scattered light images of debris
temperatures ranging from several hundred Kel- disks around HD 53143 and HD 139664. Astrophys
J 637:L57–L60
vin to approximately 40 K, respectively Kalas P et al (2007) Discovery of extreme asymmetry in
(Hillenbrand et al. 2008). For example, the Epsi- the debris disk surrounding HD 15115. Astrophys
lon Eridani debris disk can be separated into five J 661:L85–L88
components (Backman et al. 2009), and the HR Kalas P et al (2008) Optical images of an exosolar planet
25 light-years from Earth. Science 322:1345–1348
8799 disk has three components (a warm grain Kuchner MJ, Holman MJ (2003) The geometry of reso-
region interior to the three planets and two colder nant signatures in debris disks with planets. Astrophys
components exterior to the planets; Su J 588:1110–1120
616 Decarboxylation
Lagage PO, Pantin E (1994) Dust depletion in the inner there are numerous decarboxylation reactions
disk of beta Pictoris as a possible indicator of planets. which are important in biochemistry, for exam-
Nature 369:628–630
Lagrange AM, Gratdour D, Chauvin G et al (2009) ple, in the Krebs cycle, and in the biosynthesis of
A probable giant planet imaged in the beta Pictoris many neurotransmitters such as dopamine and
disk. Astron Astrophys 493:L21–L25 serotonin. It is also one of the important degrada-
Marois C, Macintosh B, Barman T et al (2008) Direct tion mechanisms of amino acids and fatty acids in
imaging of multiple planets orbiting the star HR
8799. Science 322:1348 the environment.
Ozernoy LM, Gorkavyii NN, Mather JC, Taidakova TA
(2000) Signatures of exosolar planet in dust debris
disks. Astrophys J 537:L147–L151 See Also
Rieke GH et al (2005) Decay of planetary debris disks.
Roques F, Scholl H, Sicardy B, Smith BA (1994) Is there a ▶ Amino Acid
planet around Beta Pictoris? Perturbations of a planet ▶ Carboxylic Acid
on a circumstellar disk. Icarus 108:37–58 ▶ Krebs Cycle
Smith BA, Terrile RJ (1984) A circumstellar disk around
beta Pictoris. Science 226:1421–1424
Song I et al (2003) New members of the TW Hydrae
association, beta Pictoris moving group and Tucana/
Horologium association. Astrophys J 599:342–350
Su KYL et al (2009) The debris disk around HR 8799. Decay Constant
Wyatt M (2008) Evolution of debris disks. Annu Rev
Francis Albarède
Zuckerman B (2001) Dusty circumstellar disks. Annu Rev Ecole Normale Supérieure de Lyon, Lyon,
Astron Astrophys 39:549–580 France
Zuckerman B, Song I (2004) Dusty debris disks as sign-
posts of planets: implication for spitzer space tele-
scope. Astrophys J 603:738–743
Definition
The decay constant (symbol, l and units, s1 or

a1) of a radioactive nuclide is its probability of
Decarboxylation
decay per unit time. The number of parent
nuclides P therefore decreases with time t as
dP/P dt = l. The energies involved in the
binding of protons and neutrons by the nuclear
forces are ca. 1,000,000 times stronger than those
Japan
of the electronic and molecular forces. Decay
probabilities and ls are insensitive not only to
USA
temperature and pressure but also to the strength
of the bonds in which the radioactive element is
Washington, DC, USA
held. The decay constant relates to the ▶ half-life
of the nuclide T 1/2 through T 1/2 = ln 2/l.
Definition See Also
Decarboxylation is a chemical reaction in which ▶ Earth, Age of

a carboxyl group (-COOH) is lost from a parent ▶ Geochronology
molecule as CO2. Many carboxylic acids will ▶ Half-Life
decarboxylate under even mild heating, and ▶ Radioactivity
Deep Impact 617
measured positively (+) north or negatively ()

Deccan Trapps south of the celestial equator from 0 to 90 .
Nicholas Arndt See Also

▶ Coordinate Systems
Definition ▶ Right Ascension
The Deccan ▶ Trapps (or Traps) are a volcanic D

plateau or a large igneous province located in
west-central India. They are a thick Deep Biosphere
(500–3,000 m) and large (1.5 106 km3) sequence
of basaltic lava flows that erupted subaerially and Felipe Gomez
rapidly onto the Indian continent about 67–64 Ma Centro de Astrobiologı́a (CSIC/INTA), Instituto
ago. The combined effects of large amount of sulfur Nacional de Técnica Aeroespacial, Torrejón de
gases released from these volcanic eruptions and Ardoz, Madrid, Spain
the impact of an asteroid in the Yucatan peninsula
(the ▶ Chicxulub crater) probably caused the
Cretaceous-Tertiary ▶ mass extinction that Definition
included the dinosaurs. It is generally believed
that the trapps formed through melting of a large Biosphere refers to the sum of all Earth ecosystems,
mantle plume beneath the Indian continent. in contracts to deep biosphere which refers to the
dark habitats physically located below the surface
See Also of continents and the bottom of the ocean. These
hidden environments exist in permanent darkness
▶ Basalt with no possibility of light cycle for metabolic
▶ Chicxulub Crater activity. Those habitats are poorly understood due
▶ Mantle Plume, Planetary to the technological difficulties to reach them with-
▶ Mass Extinctions out contamination and to characterize them.
▶ Trapps
Deep Impact
Declination
Hervé Cottin
Daniel Rouan Laboratoire Interuniversitaire des Systèmes
LESIA, Observatoire Paris-Site de Meudon, Atmosphériques, Université Paris Est-Créteil,
Meudon, France Créteil, France
The declination is one of the two coordinates of Deep Impact was a (NASA) mission launched on
the equatorial system of reference, used to locate January 12, 2005, made of a space probe with
a celestial object on the sky. It is the one coordi- limited instrumentation (optical imaging and
nate measuring the angular distance north or infrared spectral mapping) and carrying a
south of the celestial equator, along a meridian 370 kg impactor made mainly of copper. The
passing through that object. Declination is impactor collided with the nucleus of comet
618 Deep Subsurface Microbiology
Deep Impact, Fig. 1 Tempel 1 impact site. This pair of NASA’s Stardust spacecraft, shows the impactor erased
images shows the area affected by the impactor released that dark mound and flattened the area. The outer circle
by NASA’s Deep Impact spacecraft. On the left, the image annotated on the right-hand image shows the outer rim of
from Deep Impact spacecraft shows a dark mound about the crater, and the inner circle shows the crater floor. The
50 m in size (inside a yellow circle that shows the area hit crater is estimated to be 150 m in diameter (Credit: NASA/
by the impactor). The image on the right, obtained by JPL-Caltech/University of Maryland/Cornel)
9P/Tempel on July 4, 2005, with a relative speed which flew over the nucleus of comet 9P/Temple
around 10 km/s. The observation of the resulting on February 15, 2011 (Fig. 1).
excavation should have allowed to measure inner
properties of cometary nuclei. But, the unex-
pected large amount of dust released by
See Also
the impact prevented any direct observation of
the artificial crater. However, Deep Impact obser-
▶ Comet
vations returned unprecedented images of a com-
▶ EPOXI Mission
etary nucleus and information about the nucleus
▶ Star Dust
structure and composition. Then, the Deep Impact
spacecraft was redirected toward comet 103P/
Hartley through an extended mission. The space-
craft keeps its original name of Deep Impact,
while the next leg of the mission after the comple- Deep Subsurface Microbiology
tion of the main objective is called EPOXI. The
flyby occurred on November 4, 2010, and returned Tullis C. Onstott
high-quality pictures of this comet. After a com- Department of Geosciences, Princeton
munication failure on August 8, 2013, and several University, Princeton, NJ, USA
unsuccessful attempts to restore the contact, the
spacecraft was declared lost by NASA on Septem-
ber 19, 2013. Keywords
Follow-Up Biogeochemistry; Ecohydrology; Enzymes;

In an attempt to see the crater created formerly, Genomics; Isotope geochemistry; Microbial
NASA rerouted the Stardust-NExT spacecraft metabolism
Deep Subsurface Microbiology 619
Synonyms teams of scientists that organized and executed

these campaigns and examined the cores com-
Subsurface biota prised microbiologists, geochemists, geologists,
and hydrologists – some well seasoned and others
young and ambitious, many from DOE laborato-
Definition ries and equally many from academia, all
supported by DOE’s Subsurface Science Program
The science of deep subsurface microbiology (SSP) and led, inspired, and motivated by the pro-
involves the study of non-phototrophic microor- gram’s manager, Frank Wobber. Other groups D
ganisms, ecosystems, and biogeochemical fluxes formed quickly to explore the subsurface for life.
and cycles that occur beneath the soil zones, active Chapelle and Lovley at USGS began examining
layers, and lake beds of continents and the seafloor. Atlantic Coastal Plain aquifers at the same time
Deep subsurface ecosystems are distinct from that DOE was working near SRP. In 1986
those that inhabit most surface soils or deep ocean Whelan’s group from WHOI and, later in 1992,
environments in that they are primarily hypoxic to Parkes’ group from the Univ. of Bristol initiated
anoxic; the estimated microbial cell turnover times studies of sub-seafloor sediments. In 1992, Peder-
are on the order of decades to centuries (Phelps sen from Gothenburg Univ. published his first
et al. 1994) and the principal energy sources orig- results from a deep, fractured granite aquifer in
inate from the geosphere (Parkes et al. 2007). Like Sweden. Other groups built upon earlier Russian
surface ecosystems, the subsurface ▶ biosphere studies and began examining the fluids associated
appears to control groundwater chemistry, mineral with petroleum reservoirs. As summarized in
diagenesis, and organic degradation. Whitman et al. (1998), a substantial fraction of
the Earth’s living carbon biomass resides beneath
the seafloor and soil zones, although just how
Overview massive remains uncertain.
Although the Whitman et al. (1998) review was
In June 1986, at the Savannah River Plant (SRP), thorough at its time, the uncertainties in estimated
South Carolina, the modern era of subsurface global subsurface biomass remain large, and many
microbiology was launched with a few hundred subsurface environments had not been sampled
thousand dollars from a US Dept. of Energy and remain to this day unsampled. Studies of the
(DOE), environmental monitoring program, sav- marine deep sub-seafloor biosphere began later
ings from altering well completion designs. From than those for terrestrial settings, but they now
this auspicious beginning, three wells were cored take the lead despite sampling challenges. Both
attaining depths of 200 m using chemical and deep hot, rift basin-related subsurface marine che-
physical tracers designed by Phelps to constrain moautotrophic ecosystems and the not so hot,
microbial drilling contamination. Abundant and sub-seafloor sediments that represent vast, cold
diverse microbial communities were observed in heterotrophic ecosystems functioning on old, bur-
the subsurface aquifers, as discussed in a special ied photosynthate have been sampled. With
issue of Geomicrobiology Journal in 1989. During respect to the latter, Lipp et al. (2008) reported a
the decade that followed, a wide range of environ- correlation between the biomass and the concen-
ments and rock types were cored during seven tration of organic photosynthate in marine sedi-
field campaigns with a variety of fluids and tracers ments that constrains the global marine subsurface
mostly to depths of several 100 m. By biomass to lower values (Kallmeyer et al. 2012).
piggybacking on petroleum exploration drilling The continental deep subsurface biosphere con-
programs, microbial samples representative of tains what could be termed “warm,” <100 C
their native subsurface environment were recov- ecosystems that are increasingly chemoautotro-
ered from depths as great as 2.7 km for subsurface phic with depth. The rock-water interactions that
microbial and biogeochemical investigations. The fuel these continental subsurface ecosystems are
620 Deep Subsurface Microbiology
not dependent upon localized magmatic heat ▶ Biogeochemical Cycles

input, for example, spreading centers or hot ▶ Biosphere
spots, but instead are influenced by regional, topo- ▶ Chemolithotroph
graphically driven meteoric fluid flow. In the case ▶ Energy Conservation
of the elevated plateaus and mountain ranges ▶ Extreme Environment
groundwater can penetrate to kilometer depths ▶ Lithotroph
(Colwell et al. 1997). In the lower elevation gra- ▶ Prokaryote
nitic aquifers of the Fennoscandian Precambrian ▶ Thermophile
shield (Pedersen 1997) and Atlantic Coastal Plain
aquifers, the regional flow is slower and shallower
(Phelps et al. 1994).
Assuming a maximum temperature for life of Biddle JF, Fitz-Gibbon S, Schuster SC, Brenchley E,
120 C, mean annual surface temperatures that House CH (2008) Metagenomic signatures of the
can range from 10 to 30 C, and geothermal Peru margin subseafloor biosphere show a genetically
gradients ranging from 7 to 80 C/km, the esti- distinct environment. Proc Natl Acad Sci U S A
105:10583–10588
mated depth to the base of the deep subsurface Bidle KD, Lee SH, Marchant DR, Falkowski PG (2007)
biosphere ranges from 1 to 20 km (see Zhang Fossil genes and microbes in the oldest ice on Earth.
et al. 2005). Proc Natl Acad Sci U S A 104:13455–13460
Metagenomes have been recovered from Chivian D, Alm E, Brodie E, Culley D, Dehal P,
DeSantis T, Gihring T, Lapidus A, Lin L-H,
million-year-old ice (Bidle et al. 2007), tens of Lowry S, Moser D, Richardson P, Southam G,
millions of years old, kilometers deep, saline Wanger G, Pratt L, Andersen G, Hazen T,
fracture water (Chivian et al. 2008), and Brockman F, Arkin A, Onstott T (2008) Environmen-
subseafloor sediment environments where fluid tal genomics reveals a single species ecosystem deep
within the Earth. Science 322:275–278
flux is negligible (Biddle et al. 2008; Chivian et Colwell FS, Onstott TC, Dilwiche ME, Chandler D,
al. 2008). These studies revealed that in order to Fredrickson JK, Yao Q-J, McKinley JP, Boone DR,
repair their DNA from radiation damage, subsur- Griffiths R, Phelps TJ, Ringelberg D, White DC,
face organisms must metabolize extremely LaFreniere L, Balkwill DL, Lehman RM, Konisky J,
Long PE (1997) Microorganisms from deep, high tem-
slowly without encountering entropic death. perature, sandstones: constraints on microbial coloni-
One long-term source of renewable chemical zation. FEMS Microbiol Rev 20:425–435
energy is radiolysis of water and serpentinization D’Hondt S, Jorgensen BB, Miller DJ, Batzke A, Blake R,
of mafic and ultramafic rock, which provide Cragg BA, Cypionka H, Dickens GR, Ferdelman T,
Hinrichs KU, Holm NG, Mitterer R, Spivack A, Wang
sustained sources of oxidants and reductants to GZ, Bekins B, Engelen B, Ford K, Gettemy G, Ruth-
these terrestrial and marine subsurface chemoau- erford SD, Sass H, Skilbeck CG, Aiello IW, Guerin G,
totrophic microbial ecosystems when organic House CH, Inagaki F, Meister P, Naehr T, Niitsuma S,
carbon is scarce (Lin et al. 2005; Sherwood Lollar Parkes RJ, Schippers A, Smith DC, Teske A, Wiegel J,
Padilla CN, Acosta JLS (2004) Distributions of micro-
et al. 2014). Finally, subsurface microorganisms, bial activities in deep subseafloor sediments. Science
and even the DNA fragments of deceased subsur- 306:2216–2221
face microorganisms, may represent a pool of Edwards KJ, Bach W, McCollom TM
ancient DNA or paleome (Inagaki et al. 2005), (2005) Geomicrobiology in oceanography: microbe-
mineral interactions at and below the seafloor. Trends
and the differences between their gene sequences Microbiol 13:440–456
and those of related surface organisms may be a Inagaki F, Okada H, Tsapin AI, Nealson KH (2005) The
direct evolutionary record (Lau et al. 2014). paleome: a sedimentary genetic record of past micro-
bial communities. Astrobiology 5:141–153
Kallmeyer J, Pockalny R, Adhikaria RR, Smith DC,
D’Hondt S (2012) Global distribution of microbial
See Also abundance and biomass in subseafloor sediment. Proc
Natl Acad Sci U S A. doi:10.1073/pnas.1203849109
▶ Archaea Lau CYM, Cameron C, Magnabosco C, Brown CT,
Schilkey F, Grim S, Hendrickson S, Pullin M,
▶ Bacteria
Deep-Sea Microbiology 621
Lacrampe-Couloume G, Sherwood Lollar B, van Definition

Heerden E, Kieft TL, Onstott TC (2014) Phylogeny
and phylogeography of functional genes shared among
seven terrestrial subsurface metagenomes reveal N- The deep-sea corresponds to depths of 1,000 m
cycling and microbial evolutionary relationships. and more. It is a dark (solar light does not pene-
Front Microbiol. doi:10.3389/fmicb.2014.00531 trate beyond 100 or 200 m) and cold environment
Lin L-H, Hall JA, Lippmann J, Ward JA, Sherwood-Lollar (about 2 C), exposed to elevated hydrostatic
B, Onstott TC (2005) Radiolytic H2 in the continental
crust: nuclear power for deep subsurface microbial pressure. There is no photosynthesis and the
communities. Geochem Geophys Geosyst 6:X. sparse deep-sea organisms that inhabit this envi-
doi:10.1029/2004GC000907 ronment depend on primary production occurring D
Lipp JS, Morono Y, Inagaki F, Hinrichs K-U (2008) Sig- in the euphotic zone. However, in certain areas,
nificant contribution of archaea to extant biomass in
marine subsurface sediments. Nature 454:991–994 such as cold seep areas or hydrothermal vents,
Parkes RJ, Wellsbury P, Mather ID, Cobb SJ, Cragg BA, fluids enriched with hydrogen sulfide or methane
Hornibrook ERC, Horsfield B (2007) Temperature are discharged in deep waters. These chemicals
activation of organic matter and minerals during burial serve as energy sources for symbiotic microor-
has the potential to sustain the deep biosphere over
geological timescales. Org Geochem 38:845–852 ganisms that provide invertebrates (clams, tube
Pedersen K (1997) Microbial life in deep granitic rock. worms, shrimps) with organic matter produced
FEMS Microbiol Rev 20:399–414 by chemosynthesis.
Phelps TJ, Murphy EM, Pfiffner SM, White DC
(1994) Comparison between geochemical and biolog-
ical estimates of subsurface microbial activities.
Microb Ecol 28:335–349 History
Sherwood Lollar B, Onstott TC, Lacrampe-Coulome G,
Ballentine C (2014) Continental lithosphere doubles Deep-sea microbiology began with the report of a
global H2 flux estimates for the deep biosphere. Nature
516:379–382. doi:10.1038/nature14017 French microbiologist Certes (1884) that micro-
Whitman WB, Coleman DC, Wiebe WJ (1998) Prokary- organisms existed in waters and sediments col-
otes: the unseen majority. Proc Natl Acad Sci U S A lected at 5 km depth. A second start occurred with
95:6578–6583 the pioneer work of US microbiologists C. Zobell
Zhang G, Dong H, Xu Z, Zhao D, Zhang C (2005)
Microbial diversity in ultra-high-pressure rocks and and R. Morita in the 1950s. The availability of
fluids from the Chinese continental scientific pressure-retaining sampling devices during the
drilling project in China. App Env Microbiol 1960s and 1970s of manned and robotic under-
71:3213–3227 water vehicles renewed deep-sea microbiology,
and several dedicated microbiology teams carried
out extensive research particularly in the USA,
Europe (Germany, France, the UK), and Japan.
Deep-Sea Microbiology
Daniel Prieur Overview

Université de Bretagne Occidentale (University
of Western Britanny), Brest, France For the early scientists involved in deep-sea
Institut Universitaire Européen de la Mer microbiology, the first questions concerned the
(IUEM), Technopôle Brest–Iroise, Plouzané, abundance and metabolic activities of bacteria
France under in situ conditions (cold temperature,
oligotrophy, and elevated pressures). The first
experiments concluded that abundant (106 cells
Keywords ml1 of deep water, 109 cells g1 of deep sedi-
ment) deep-sea bacteria were not very active
Archaea; Bacteria; Diversity; Extremophile; under in situ conditions, essentially because of
Hydrostatic pressure; Metabolic activity; elevated pressures. This view totally changed
Piezophiles with the discovery of piezophilic bacteria living
622 Degassing
in the digestive tracts of deep-sea invertebrates. See Also

In the last decades, many piezophilic deep-sea
bacteria have been isolated and described: all ▶ Archaea
are heterotrophic, aerobes, or facultative anaer- ▶ Barophile
obes and belong to the phylum of ▶ Black Smoker
▶ Proteobacteria. More recently, ▶ Archaea ▶ Deep Subsurface Microbiology
also have been reported from molecular studies, ▶ Extremophiles
and although their abundance is about 104 cells ▶ Hot Vent Microbiology
ml1 in the deep layers, almost nothing is known ▶ Hydrothermal Vent Origin of Life Models
about their physiology and metabolism. The dis- ▶ Hyperthermophile
covery of hydrothermal vents in 1977 has consid- ▶ Piezophile
erably changed our views about deep-sea biology. ▶ Psychrophile
At vents and cold seeps, animal communities are
dense due to their symbiotic associations with
chemolithoautotrophic bacteria that obtain their
energy from the oxidation of reduced compounds Bartlett DH (1999) Microbial adaptations to the
(hydrogen sulfide, methane) present in emitted psychrosphere/piezosphere. J Mol Microbiol
fluids and provide invertebrates with this Biotechnol 1:93–100
chemosynthetically produced organic matter. Certes A (1884) Sur la culture, à l’abri des germes
atmosphériques, des eaux et des sediments
Also, various hyperthermophilic Bacteria and rapports par les expeditions du Travailleur et du
Archaea thrive in the black smokers that flow out Talisman, 18821883. C R Acad Sci Paris
of deep-sea vents. Among them are the most 98:690–693
impressive extremophiles living on Earth, such Corliss JB et al (1979) Submarine thermal springs on the
galapagos rift. Science 203:1073–1083
as the anaerobic archaea Pyrolobus fumarii, Jannasch HW, Taylor CD (1984) Deep-sea microbiology.
which grows optimally at 106 C (maximum Annu Rev Microbiol 38:487–514
113 C), or Pyrococcus yayanosii, which grows Prieur D, Marteinsson VT (1998) Prokaryotes living under
optimally at 95 C with an optimal pressure of elevated hydrostatic pressure. Adv Biochem Eng
Biotechnol 61:23–35
52 MPa (maximum 120 MPa) and cannot grow Prieur D et al (2010) Piezophilic prokaryotes. In: Sebert
under atmospheric pressure. Both organisms resist P (ed) Comparative high pressure biology.
normally sterilizing conditions in an autoclave: Science Publishers/Jersey, Enfield/Plymouth,
they have survived for more than 24 h at 120 C pp 285–322
Yayanos AA (1986) Evolutional and ecological
and up to 160 MPa for P. yayanosii). implications of the properties of deep-sea
During the last 20 years, deep-sea microbiolo- barophilic bacteria. Proc Natl Acad Sci U S A
gists have investigated an unexpected ecosystem: 83:9542–9546
the deep biosphere, a term used to describe the
enormous bacterial mass in the upper kilometers
of the Earth’s crust. J. Parkes first reported the
existence of living bacterial cells in deep marine
sediments, several hundred meters below the sea- Degassing
floor. They were recently reported at a depth of
1,600 m in rocks deposited about 110 million Francis Albarède
years ago. These ecosystems represent the largest Ecole Normale Supérieure de Lyon, Lyon,
habitat for ▶ prokaryotes on Earth. Some organ- France
isms of the deep biosphere might be totally inde-
pendent from solar energy (using abiotically
produced carbon and energy sources). A parallel Synonyms
might exist between the emergence of the first life
and the deepest parts of the biosphere. Outgassing
Delta, Isotopic 623
Definition ▶ Mars with an average distance of 23,458 km

to the center of the ▶ planet and an orbital period
The term “degassing” (or outgassing) refers to the of 1.26 days (IAU 2006). Deimos has an overall
process of releasing atmospheric gas present in smooth surface and measures 15.0 12.2
the mantle or in magmas when its amount 10.4 km; its bulk density is about 1.5 g/cm3.
exceeds the solubility limit. The distribution of The heavily cratered surface indicates an old
volatile elements, such as carbon dioxide (CO2) formation age. Two hypotheses are actually
or argon (Ar), between a solid (or a liquid) and a discussed to explain the origin of Deimos: cap-
gas, reaches equilibrium when further transfer ture of a D-type ▶ asteroid or reaccretion of Mar- D
from one phase to another entails an overall tian ejecta.
increase of the system energy. Henry’s law states
that, at equilibrium, the amount of volatile ele-
ment dissolved is proportional to its partial pres- See Also
sure in the gas phase at a specific temperature.
Outgassing of planetary mantles by volcanic ▶ Asteroid
activity is viewed as a major pathway to the ▶ Mars
formation of secondary atmospheres on telluric ▶ Phobos
planets. ▶ Planet
See Also
▶ Earth’s Atmosphere, Origin and Evolution of

▶ Earth, Formation and Early Evolution Delta, Isotopic
▶ Mantle Volatiles
▶ Noble Gases Daniele L. Pinti
Dehydration
Definition
▶ Desiccation
The delta notation (symbol: d) expresses the var-
iation of the isotopic ratio of an element R (e.g.,
d18 O ¼ 18 O=16 O), relative to the isotopic ratio of
Deimos a standard Rstd (e.g., d18 OVSMOW ¼ 18 O=16 O ¼
2005:20 0:45 106 , where V-SMOW is
Harald Hoffmann Standard Mean Ocean Water). The delta notation
DLR, Institute of Planetary Research, Berlin, is usually expressed in terms of per mil (parts
Germany per thousand, ‰). The delta notation was
invented first to express in a convenient and read-
able way the variability of isotopic ratio in many
Definition natural systems in which the range is in the third
to fifth decimal place. It also removes the need to
Deimos is the outer of the two Martian satellites report absolute isotope abundances and replaces
that were discovered by Asaph Hall in 1877. It is them with relative deviations with respect to
in nearly circular synchronous orbits around well-distributed standards.
624 Denaturation
See Also See Also
▶ Carbon Isotopes as a Geochemical Tracer ▶ Nucleic Acids

▶ Hydrogen Isotopes ▶ Polymer
▶ Iron Isotopes ▶ Protein
▶ Nitrogen Isotopes
▶ Oxygen Isotopes
▶ Silicon Isotopes
▶ Sulfur Isotopes
Denitrification
Ricardo Amils
Denaturation Universidad Autónoma de Madrid, Madrid,
Spain
Synonyms
Japan
Nitrate reduction
USA
Definition
Washington, DC, USA
Denitrification is a microbial ▶ anaerobic respira-
tion in which oxidized nitrogen compounds
(e.g., NO 3 ) are used as ▶ electron acceptors,
which may ultimately produce molecular nitrogen
Definition (N2) through a series of intermediate gaseous nitro-
gen oxide products. Inorganic nitrogen compounds
In biochemistry, denaturation is the process by are some of the most common electron acceptors
which ▶ proteins or nucleic acids lose their used in anaerobic respiration. One of the most
native quaternary, tertiary, and/or secondary common electron acceptors for anaerobic respira-
structure. Denaturation can be caused by condition is nitrate, which can be reduced to NO2 , N2O,
tions such as strong acid or base, concentrated NO, and N2. The process is the main means by
salt, organic solvents, or heat. Protein which N2 is formed biologically. Denitrification
denaturation generally results in the loss of completes the nitrogen cycle, by returning N2 to
enzymatic activity. Denatured proteins may the atmosphere. The process is performed primar-
become insoluble or aggregate. ▶ Nucleic acid ily by heterotrophic bacteria (such as Pseudomo-
denaturation results in the separation of nads), although autotrophic denitrifiers have also
hydrogen-bonded complementary regions been identified (e.g., Thiobacillus denitrificans).
when the hydrogen bonds between the strands Generally, several species of bacteria are involved
are broken. These bonds may be restored by in the complete ▶ reduction of nitrate to molecular
annealing. Denaturation typically does not result nitrogen, and more than one enzymatic pathway
in the breaking of the covalent bonds which hold has been identified in the reduction process. Deni-
a ▶ polymer together. Denaturation is often trification is a widespread transduction system
reversible in proteins and nucleic acids. throughout the phylogenetic tree.
Dense Core 625
See Also
Dense Core
▶ Electron Acceptor Steven W. Stahler
▶ Energy Conservation Department of Astronomy, University of
▶ Heterotroph California, Berkeley, CA, USA
▶ Nitrogen Cycle, Biological
▶ Reduction
Keywords D
Star formation
Dense Cloud
Definition
William M. Irvine
Dense cores are the smallest and densest types of
molecular clouds. They are particularly signifi-
cant as being the birthplaces of both single and
binary stars. A large fraction of dense cores, in
Definition
fact, already contain young stars. Many of the rest
exhibit a slow, inward contraction, which could
In relation to the ▶ interstellar medium, the term
be the prelude to the full collapse that results in
dense cloud is only vaguely defined. It is some-
star formation. Both types of dense cores have
times used to refer to that portion of ▶ molecular
temperatures near 10 K. They are slowly rotating
clouds with densities exceeding a few 100 atoms
and appear elliptical when seen in projection onto
per cubic centimeter. Some authors use dense to
the sky. In three-dimensional space, they are pro-
refer to “dense cores,” those regions in molecular
late spheroids or perhaps more complex, triaxial
clouds with densities of 105 atoms per cc or
configurations.
greater, where stars may form or are
forming. The gas phase of a dense interstellar
cloud consists almost entirely of molecular
Overview
hydrogen (H2), with an admixture of ▶ interstel-
lar dust. Since molecular hydrogen lacks an elec-
The Milky Way, like other spiral galaxies,
tric dipole moment and hence has no allowed
contains both stars and more rarefied, interstellar
rotational transitions, the gas in dense clouds is
gas. About half the mass of this material is in
usually traced by observing the millimeter-
molecular clouds, so called because they are
wavelength rotational transitions of CO, the
chiefly made of hydrogen molecules. Molecular
second most abundant gas phase constituent
clouds have a wide range of sizes and
(typically about 1 part in 10,000 by number,
masses. They are not uniform, but contain
relative to H2).
nested substructures. The smallest coherent
structures, about 0.1 pc in size and with
masses a few times the solar value, are known
See Also as dense cores. They are of special importance,
since they produce new stars. It is now believed
▶ Interstellar Dust that most binary stars are also created within
▶ Interstellar Medium dense cores.
626 Deoxyribonucleic Acid
Throughout most of the volume of molecular References and Further Reading

clouds, the gas has supersonic internal motion.
The dynamical pressure associated with this Jones CE, Basu S (2002) The intrinsic shapes of molecular
cloud fragments over a range of length scales.
motion helps to support the cloud for long periods
Astrophys J 569:280
of time against the force of self-gravity. The Myers PC, Evans NJ, Ohashi N (2000) Observations of
motion within dense cores, however, is infall in star-forming regions. In: Mannings V,
subsonic. Thus, these entities are in balance Boss AP, Russell SS (eds) Protostars
and planets IV. University of Arizona Press, Tucson,
between internal thermal pressure and self-
p 217
gravity. They also contain a magnetic field, Ryden BS (1996) The shapes of dense cores and bok
which aids in the support. The temperature of globules. Astrophys J 471:822
dense cores, like that of all molecular clouds, is
very low, roughly 10 K. It is the combination of
low temperature and relatively high density that
renders dense cores susceptible to collapse and
star formation. Deoxyribonucleic Acid
Almost half of observed dense cores already
enclose young stars. Since the cores contain solid ▶ DNA
dust grains along with gas, the embedded stars are
highly obscured and can only be seen at infrared
and longer wavelengths. The majority of dense
cores have no internal stars. The bulk properties
of these barren cores are similar to those Deoxyribose
containing stars. However, a high fraction of star-
less cores are undergoing inward contraction, as Henderson James (Jim) Cleaves II
seen by Doppler-shifted emission lines from var- Earth–Life Science Institute (ELSI), Tokyo
ious trace molecules. This motion, which is sub- Institute of Technology, Meguro–ku, Tokyo,
sonic, could be the prelude to the eventual Japan
collapse that forms a star or binary near the Institute for Advanced Study, Princeton, NJ,
dense core’s center. USA
Dense cores appear to have elliptical contours, Blue Marble Space Institute of Science,
when they are mapped in molecular lines. Each Washington, DC, USA
ellipse is a projection onto the sky of the objects’ Center for Chemical Evolution, Georgia Institute
true, three-dimensional shape. This true shape of Technology, Atlanta, GA, USA
has been difficult to ascertain, with some
researchers favoring prolate spheroids (stubby
cigars) and others more complicated, triaxial con- Definition
figurations. All agree, however, that the dense
cores’ shapes do not stem from their rotation. Deoxyribose is an aldopentose (C5H10O4) found
The latter has been observed, and the rate is too in the backbone of DNA. It exists in two stereo-
small to cause significant distortion of the cloud. isomeric forms, D and L. The isomer used in
DNA is the D-form. It is derived biologically
from D-ribose by reduction of the C2 hydroxyl
See Also group via a radical mechanism. It is typically
found in deoxynucleotides in the pentofuranose
▶ Fragmentation of Interstellar Clouds form. In DNA, it is linked via the 30 and 50
▶ Gravitational Collapse, Stellar hydroxyl groups to connecting phosphate groups.
▶ Molecular Cloud When attached to a nitrogenous base, it becomes
▶ Protostars part of a deoxynucleoside.
Desiccation 627
See Also per billion) by its characteristic fluorescence at an

optimal excitation wavelength. In the case of
▶ Aldose carboxylic acids, the reagents are usually alco-
▶ Carbohydrate hols in the presence of catalysts that convert the
▶ Nucleoside original molecules into esters for GC analyses.
▶ Nucleotide For the GC determination of amino acids, alco-
hols in the presence of catalysts are used to
first convert the carboxyl groups to esters and
reagents such as trifluoroacetic anhydride then D
used to add components to the primary amino
Depyrogenation
groups to prevent their ionization. For the fluo-
rescent analyses of amino acids, several reagents,
Catharine A. Conley
such as fluorescamine or ortho-phthalaldehyde
(OPA), are added to the primary amino group
producing highly fluorescent products that can
be analyzed using high-performance liquid
Definition
chromatography.
A pyrogen is a substance which causes, when in
the blood flow of a homeothermic animal, a rise in
the body or central temperature. The pyrogens can
See Also
be synthetic molecules but often are toxins pro-
duced by pathogenic bacteria. Depyrogenation is
▶ Amino Acid
the process of removing these pyrogens from a
▶ Chromatography
solution or compound. Depyrogenation has to be
▶ Fluorescence
performed on containers and reactants to be used
for some life detection tests.
Derivatization Desiccation
Jeffrey Bada Gerda Horneck

Scripps Institution of Oceanography, La Jolla, DLR German Aerospace Center, Institute of
CA, USA Aerospace Medicine, Radiation Biology, Köln,
Germany
Definition
Keywords
Derivatization is the process by which an organic
Anhydrobiosis; Cryptobiosis; Desiccation;
compound is reacted with a reagent, or reagents,
Dehydration; Habitability of Mars; Survival
to yield products suitable for analyses by various
techniques. In many cases, the products are either
a volatile adduct suitable for analysis by ▶ gas
chromatography (GC) or a highly fluorescent Synonyms
product that can be separated by liquid ▶ chro-
matography and detected at very low levels (parts Dehydration; Desiccation effects
628 Desiccation
Definition high radiation resistance and likewise a high

resistance to desiccation. It has been suggested
Desiccation describes a process which causes that the high radiation resistance is an evolution-
loss of liquid water from a solid surface or from ary consequence of an earlier ▶ adaptation to
a porous medium (soil, food, ecosystems, indi- prolonged periods of cellular desiccation
vidual organisms, etc.). This process leads to (Mattimore and Battista 1996).
drying up of the matter. In biology, several organ- From the viewpoint of astrobiology, the ▶ sur-
isms have developed strategies to tolerate desic- vival of microorganisms in outer space is of
cation imposed on them. interest, where they are exposed to a complex
matrix of parameters, including the extremely
dehydrating space vacuum. Space exposure
Overview experiments have shown that although a variety
of microorganisms are known to be desiccation
Water is one of the essentials for all forms of life resistant, not many were able to cope with the
on the Earth. Dehydration causes severe damage mechanical stress of the high vacuum of space.
to various cell components: Lipid membranes The record of survival in space was reached by
may change from planar bilayers to cylindrical Bacillus subtilis spores, of which 70–90 % sur-
bilayers, and carbohydrates, proteins, and nucleic vived after 6 years in space vacuum (Horneck
acids may undergo amino-carbonyl reactions et al. 2010).
(Maillard reactions) that result in cross-linking ▶ Mars is an important target of astrobiology,
and finally polymerization of the biomolecules where the assessment of its habitability largely
(Cox 1991, 1993). These structural changes can depends on the availability of liquid water.
give rise to functional changes, such as inhibited Because Mars is cold, but not always, and
or altered enzyme activity, changes in membrane extremely dry, but perhaps not everywhere, the
permeability, and alteration of genetic informa- concept of a special region on Mars was devel-
tion. The latter change is especially dramatic, oped as a way to refer to those places where the
because it may lead to cell death or mutagenesis conditions on Mars might be conducive to micro-
(Munakata et al. 1997). bial growth (Kminek et al. 2010). It was con-
Certain organisms possess the ability to with- cluded that any region experiencing
stand extreme dryness, such as plants and animals temperatures >25 C for a few hours a year
living in arid or periodically arid environments and a ▶ water activity >0.5 can potentially
(temporary rivers or ponds), where they may face allow the replication of terrestrial microorgan-
the challenge of desiccation. In response to such isms. Such special regions on Mars require spe-
extreme and unfavorable environmental condi- cial ▶ planetary protection measures for landers
tions, organisms may resume a cryptobiotic and rovers sent to the surface of Mars (COSPAR
state, which refers to a state of the organism that Planetary Protection Policy: http://cosparhq.
has no detectable metabolic activity. cnes.fr/Scistr/PPPolicy%2820-July-08%29.pdf).
Anhydrobiosis refers to the state of an organism In biotechnology, freeze-drying or lyophiliza-
that survives the loss of (almost) all body water. tion is used as a method of preserving microor-
In highly desiccation-resistant bacterial spores, ganisms or tissue without destroying their
the water content in the ▶ spore core is naturally physical structure and viability (Morgan and
reduced to 25–45 % of their wet weight. As a Vesey 2009).
consequence, proteins are immobile and enzymes
inactive during the spore phase (Setlow 1994).
This low water content is one of the reasons for See Also
the high resistance of bacterial spores to a variety
of physical and chemical harmful agents. The ▶ Adaptation
bacterium Deinococcus radiodurans shows a ▶ Aerobiology
Deuterium 629
▶ Mars Deuterium
▶ Microorganism
▶ Planetary Protection Mark Dörr
▶ Space Vacuum Effects University of Southern Denmark, Odense M,
▶ Spore Denmark
▶ Survival
▶ Water Activity
Keywords D
Hydrogen; Isotope
Cox CS (1991) Roles of Maillard reactions in diseases.
Her Majesty’s Stationery Office (HMSO), London
Cox CS (1993) Roles of water molecules in bacteria and Synonyms
viruses. Orig Life Evol Biosph 23:29–36
Horneck G, Klaus DM, Mancinelli RL (2010) Space
microbiology. Microbiol Mol Biol Rev 74:121–156 Diplogen; Heavy hydrogen
Kminek G et al (2010) Report of the COSPAR mars
special regions colloquium. Adv Space Res
Mattimore V, Battista JR (1996) Radioresistance of
Deinococcus radiodurans: functions necessary to sur-
vive ionizing radiation are also necessary to survive Deuterium (Symbol: 2H, earlier, D) (from the
prolonged desiccation. J Bacteriol 178:633–637 Greek: deύteroB (deúteros) = “the second”) is
Morgan C, Vesey G (2009) Freeze-drying of microorgan- a stable ▶ isotope of ▶ hydrogen with a relative
isms. Encyclopedia of microbiology, 3rd edn.
Elsevier, New York, pp 162–173 atomic mass of 2.01410222 mu and a nuclear spin
Munakata N, Saito M, Takahashi N, Hieda K, Morihoshi F of 1. Mass spectrometry and various scattering
(1997) Induction of unique tandem-base change muta- experiments reveal two elementary particles in its
tions in bacterial spores exposed to extreme dryness. atomic nucleus, a positively charged proton and an
Mutat Res 390:189–195
Setlow P (1994) Mechanisms which contribute to the uncharged neutron. These observations gave rise to
long-term survival of spores of Bacillus species. its name. Other natural isotopes of hydrogen are
J Appl Bacteriol Symp 76:129S–134S Protium (1H) (one proton, 1.00782519 mu, nuclear
spin ½) and Tritium (3H) (one proton, two neutrons,
3.01610497 mu, nuclear spin ½, radioactive)
(Greenwood and Earnshaw 1984). Because of its
Desiccation Effects mass, deuterium is also called “heavy hydrogen.”
▶ Desiccation
History
Deuterium was discovered in 1931 by Harold

Detergent C. Urey, who received the Nobel Prize in chem-
istry (1934) for this finding.
▶ Amphiphile
Overview
Deuterated Ammonium Ion Hydrogen is the most abundant element in the

universe (ca. 76 % by mass). The average
▶ Ammonium, Deuterated deuterium-to-hydrogen ratio in the universe is
630 Deuterium/Hydrogen Ratio
estimated to be ca. 20 ppm (Lellouch et al. 2001). Hydrogen isotopic fractionation is very pro-
Deuterium has a natural abundance in the Earth’s nounced in cold interstellar clouds, so that the
crust of 2H/1H = 145 ppm. In the ocean, this abundance ratio of (e.g.,) DCN/HCN can be
ratio is 155.76 0.1 ppm (a ratio of 1 part per much larger than the cosmic D/H ratio. In fact,
approximately 6,420 parts) according to the doubly deuterated and even triply deuterated
Vienna Standard Mean Ocean Water (VSMOW) molecules are observed in the ▶ interstellar
(American Society of Limnology and Oceanog- medium, for example, D2CO and ND3.
raphy Inc 1995). Deuterium abundance in Jupi-
ter’s atmosphere is about 22.5 ppm (ca. 15 % of
the terrestrial deuterium-to-hydrogen ratio); that See Also
of Saturn’s atmosphere is about 17 ppm
(Lellouch et al. 2001). The Jupiter ratio presum- ▶ Deuterium/Hydrogen Ratio
ably reflects the early solar nebular ratio as well ▶ Hydrogen
as the one present during the early stages of the ▶ Isotope
universe (Lellouch et al. 2001). Most of the deu- ▶ Urey’s Conception of Origins of Life
terium in the universe is believed to have been
produced during the nucleosynthesis era of the
Big Bang (~10 s after t0), since other sources,
such as rare cluster decays, and occasional American Society of Limnology and Oceanography Inc
absorption of naturally occurring neutrons by (1995) Limnol Oceanogr 40(6):1182. http://www.aslo.
hydrogen do not account for the amount presently org/lo/toc/vol_40/issue_6/1182.pdf
observed. If the de novo production rate of deu- Greenwood N, Earnshaw A (1984) Chemistry of the ele-
ments. Pergamon Press, Elmsford
terium is very low, the amount of deuterium in Lellouch E, Bézard B, Fouchet T, Feuchtgruber H,
the universe declines steadily since it is con- Encrenaz T, De Graauw T (2001) The deuterium abun-
sumed in nuclear fusion reactions in stars. These dance in Jupiter and Saturn from ISO-SWS observa-
fusion reactions are much faster than the proton- tions. Astron Astrophys 670:610–622. doi:10.1051/
0004-6361:20010259
proton reactions that generate deuterium at very
high temperatures in stars. Gamma radiation
from ordinary nuclear fusion dissociates deute-
rium into protons and neutrons. Both reaction
kinetics and gamma radiation make it improbable Deuterium/Hydrogen Ratio
that the concentration of deuterium in the interior
of the Sun and other stars is higher than the Daniele L. Pinti
average universal abundance. In the outer solar GEOTOP Research Center for Geochemistry and
atmosphere, however, deuterium is present in the Geodynamics, Université du Québec à Montréal,
same concentration as the one estimated on Jupi- Montréal, QC, Canada
ter. The natural deuterium abundance seems to be
very similar wherever hydrogen is found.
Deuterium occurs in trace amounts naturally Synonyms
as deuterium gas (2H2 = D2), but most of the
deuterium in the universe is bound to a 1H atom, D/H
forming a gas called hydrogen deuteride (1H2H or
HD). Deuterium exchanges rapidly with hydro-
gen in many compounds, like R-O-D and R-N-D, Definition
but slowly in many R-C-D compounds
(R denotes a residue). This is the basis of impor- The deuterium/hydrogen ratio or D/H is the ratio of
tant kinetic and mechanistic measurements in the ▶ hydrogen isotopes of mass 2 (21H or D, deu-
chemical reactions. terium) and mass 1 (11H or protium). The ratio is a
Dharwar Craton 631
valuable tracer of fractionation processes in induced hydrothermal deposits, as well as

hydrogen-bearing (water, hydrocarbon) systems. It intracrater paleolake deposits, alteration min-
has been measured in hydrogenated molecules and erals, valleys and gullies, and polygonal ground.
hydrous minerals preserved in ▶ meteorites and In addition, the biological characteristics
▶ comets and compared to the ratio in the Earth’s (e.g., endolithic and extremophilic microorgan-
oceans and in hydrous minerals from mare basalts isms, little to no vegetation, low nutrients, and
and highland rocks of the ▶ Moon for determining organic biomarkers in impacted rocks) are of
the carrier of water on the Earth-Moon system. particular interest to astrobiologists.
The D/H ratio shows a heliocentric distribution D
within the ▶ Solar System, with the lowest value
See Also
measured in the Sun ( D=H ¼ 21 5 106 )
and the highest in the HCN compounds in comets
▶ Antarctica
( D=H ¼ 2300 400 106 ). This variance is
▶ Crater, Impact
mostly related to the deuterium isotopic exchange
▶ Cryosphere
rate between molecular hydrogen and water.
▶ Extremophiles
See Also ▶ Impact Melt Rock
▶ Impactite
▶ Carbonaceous Chondrite ▶ Mars Analogue Sites
▶ Chondrite ▶ Permafrost
▶ Comet ▶ Terrestrial Analog
▶ Hydrogen Isotopes
▶ Oceans, Origin of
▶ System Solar Formation, Chronology of
▶ Water, Delivery to Earth
Dewar Flask
▶ Cryostat
Devon Island
Richard Léveillé DHA

Natural Resource Sciences, McGill University,
St. Anne de Bellevue, Québec, Canada ▶ Dihydroxyacetone
Definition
Dharwar Craton
Devon Island (federal territory of Nunavut) is a
large (66,800 km2), uninhabited island of the Jean-François Moyen
Canadian Arctic Archipelago. Situated in a LMV-TL, Université Jean-Monnet,
polar desert, Devon Island is considered to be an Saint-Etienne, France
analog to ▶ Mars because it is cold, dry, and
largely non-vegetated, with a variety of analo-
gous geological and geomorphologic features. Keywords
These include the well-exposed and preserved
Haughton impact crater and associated impact- Archean; Craton; Geological province; Accretion
632 Dharwar Craton
Synonyms The western Dharwar craton is dominated by

Mesoarchean rocks, including a ca. 3.3–3.0 Ga
South Indian shield basement of TTG gneisses (the “Peninsular
Gneisses”) and two successions of greenstone
belts: an older, ca. 3.3 Ga Sargur Group and a
Definition younger, 3.0–2.7 Ga Dharwar Group unconform-
ably overlying the Peninsular Gneisses
The Dharwar craton is a part of southern India, (Chadwick et al. 2000 and references therein). It
underlain by Archean rocks and extending is further intruded by several late Archean,
roughly between the cities of Goa, Hyderabad, ca. 2.61 Ga, plutons (Jayananda et al. 2006) of
Chennai, and Mangalore in Karnataka and potassic granitoids. In contrast, the eastern
Andhra Pradesh states. Dharwar craton does contain only rare
Mesoarchean remains; it is mostly made of
Neoarchean juvenile granitoids that were
Overview emplaced ca. 2.55 Ga ago, concomitant with the
main deformation phase (Jayananda et al. 2000;
The Dharwar craton is made of orthogneisses and Chardon et al. 2002). However, recent fieldwork,
greenstone belts, collectively formed between 3.3 mapping, and geochronology allowed to further
and 2.6 Ga (Chadwick et al. 2000). It acquired its subdivide the eastern Dharwar craton and iden-
final structure between 2.56 and 2.51 Ga during a tify a third, intervening domain with distinct fea-
tectonic and metamorphic event that overprinted tures (Moyen et al. 2003; Chardon et al. 2011).
most previous structures and lithologies Greenstone belts of the eastern Dharwar craton
(Chardon and Jayananda 2008). This event was are 2.7–2.5 Ga old. The boundary between these
accompanied by the intrusion of voluminous domains has been interpreted as a tectonic suture,
granitoids (Chardon et al. 2011). Due to differen- although it lacks many typical features of modern
tial erosion and/or tilting, different crustal levels boundaries such as thrust-and-fold structures,
are exposed from south (granulite facies) to north HP/LT metamorphism, or ophiolites, and exem-
(lower grade), allowing to reconstruct a three- plifies the ambiguity of the Archean geological
dimensional section through the late Archean record.
crust. The Dharwar craton has been pervasively
To the south, the Dharwar craton is cut by affected by a ca. 2.55–2.51 Ga tectono-
younger (Proterozoic) tectonic structures; vari- metamorphic event, involving (1) the emplace-
ous granulitic domains rework the Archean ment of large volumes of mostly granitoids in the
crust. It has also been suggested (Collins eastern Dharwar, (2) high-grade (granulite facies)
et al. 2014) that the northwestern corner of the metamorphism in the lower crust now exposed in
Dharwar craton (Goa area) represents part of the southern part of the craton (these granulites
western Gondwana which docked in the either correspond to modified old gneisses or to
Neoproterozoic against the Dharwar craton juvenile magmas emplaced in the lower crust),
proper. The craton is covered by younger units and (3) deformation represented by E-W bulk
to the north and east, including Proterozoic sed- shortening of the crust, accommodated by
iments of the Cuddapah basin and flood basalts of transcurrent shear zones and (in the western
the Deccan traps. Fragments of crust of Archean Dharwar) and sinking of the dense greenstone
age are also found in the heavily reworked gran- belts into the lighter gneissic basement
ulite terrain of Peninsular India, south of the (“sagduction”) (Chardon et al. 1996).
craton (Collins et al. 2014). The Dharwar craton has been the focus of
The Dharwar craton is traditionally separated research for ancient traces of life, particularly in
into two main geological domains (west and east) the western Dharwar craton which contains
with contrasting ages and geological features. Archean sedimentary rocks, among others
Dharwar Craton
633
Dharwar Craton, Fig. 1 a) Main crustal units forming the Indian shield. b) Simplified geological map of the Dharwar craton (Modified after Chardon et al. (2008))
D
634 DHMR
several units of banded iron formations (BIFs) Ma) juvenile magmatism in the Eastern Dharwar cra-
(Manikyamba et al. 1993). In BIFs, Orberger ton, southern India: constraints from geochronology,
Nd-Sr isotopes and whole rock geochemistry. Precam-
et al. (2012) found spherules containing carbona- brian Res 99:225–254
ceous matter which might be derived from Jayananda M, Chardon DH, Peucat JJ, Capdevila R (2006)
ancient microbial activity (Fig. 1). 2.61 Ga potassic granites and crustal reworking in the
Western Dharwar craton, Southern India: tectonic,
geochronologic and geochemical constraints. Precam-
Manikyamba C, Balaram V, Naqvi SM (1993) Geochem-
See Also
ical signatures of polygenetic origin of a banded iron
formation (BIF) of the Archaean Sandur greenstone
▶ Archean Eon belt (schist belt) Karnataka nucleus, India. Precam-
▶ Archean Tectonics brian Res 61:137–164
Moyen JF, Martin H, Jayananda M (2003) Magmatism dur-
ing the accretion of the late Archaean Dharwar Craton
▶ Craton (South India): sanukitoids and related rocks in their geo-
▶ Sagduction logical context. EGS-AGU-EUG joint assembly, Nice
▶ Shield Orberger B, Wagner C, Wirth R, Quirico E, Gallien JP,
Derré C, Montagnac G, Noret A, Jayananda M,
▶ TTG
Massault M, Rouchon V (2012) Origin of iron oxide
spherules in the banded iron formation of the
Bababudan Group, Dharwar Craton Southern India.
References and Further Reading J Asian Earth Sci 52:31–42
Chadwick B, Vasudev VN, Hegde GV (2000) The

Dharwar Craton, southern India, interpreted as the
result of late Archaean oblique convergence. Precam-
brian Res 99:91–111 DHMR
Chardon, D, Jayananda M (2008) Three-dimensional field
perspective on deformation, flow, and growth of the Catharine A. Conley
lower continental crust (Dharwar craton, India). Tec-
tonics 27:TC1014 NASA Headquarters, Washington, DC, USA
Chardon DH, Choukroune P, Jayananda M (1996) Strain
patterns, decollement and incipient sagducted green-
stone terrains in the Archaean Dharwar craton (South Definition
India). J Struct Geol 18:991–1004
Chardon DH, Peucat JJ, Jayananda M, Choukroune P,
Fanning CM (2002) Archean granite-greenstone tec- Dry heat microbial reduction (DHMR) is a pro-
tonics at Kolar (South India): interplay of diapirism cess for ▶ bioburden reduction. This process con-
and bulk inhomogeneous contraction during juvenile sists of heating a piece of hardware in an oven for
magmatic accretion. Tectonics 21:7-1–7-17,
TC901032 a time depending on the set temperature. It was
Chardon D, Jayananda M, Chetty TRK, Peucat JJ developed for entire spacecrafts by ▶ NASA’s
(2008) Precambrian continental strain and shear zone Viking Project. In this case, the spacecrafts were
patterns: South Indian case. J Geophys Res Solid Earth placed for 30 h at 111.7 C. The process is
113(B8)
Chardon D, Jayananda M, Peucat JJ (2011) Lateral approved by both NASA and ESA for use with
constrictional flow of hot orogenic crust: insights spacecraft hardware over a specified range of
from the Neoarchean of south India, geological and temperature and humidity.
geophysical implications for orogenic plateaux. Geoch
Geophy Geosyst 12: GC003398
Collins AS, Clark C, Plavsa D (2014) Peninsula India in
Gondwana: the tectonothermal evolution of the south- See Also
ern granulite terrane and its Gondwana counterparts.
Gondwana Res 25:190–203 ▶ Bioburden
Jayananda M, Moyen J-F, Martin H, Peucat J-J, Auvray B,
Mahabaleswar B (2000) Late Archaean (2550–2520 ▶ Bioburden Reduction
Diagenesis 635
▶ Pasteurization
▶ Planetary Protection Diagenesis
▶ Sterilization
Nicholas Arndt
Diacetylene
Definition
D
William M. Irvine
Diagenesis refers to all physical, chemical, and
biological changes that accompany the first stage
of lithification, exclusive of surface alteration
(weathering) and metamorphism. Diagenesis is
Synonyms
the agent that transforms unconsolidated sedi-
ment into solid rock (▶ sedimentary rock)
Butadiyne; C4H2; HC4H
through a combination of compaction, recrystal-
lization, dehydration, replacement, dissolution,
and cementation processes. Diagenesis virtually
Definition
begins when a detrital grain is sedimented and
begins its interaction with the pore waters and
The molecule diacetylene is the simplest
overburden. Diagenesis ends when these pro-
polyyne, which is a hydrocarbon with alternating
cesses grade into metamorphism, and sedimen-
single and triple bonds; thus, diacetylene has the
tary textures and structures (such as bedding,
formula HCC-CCH. Diacetylene is found as a
grading, bioturbation, etc.) are lost.
minor component in the atmospheres of the
Diagenetic processes thus usually take place at
▶ giant planets, including Jupiter, Saturn, and
low temperature (<200 C) and pressure
Neptune, and in that of Saturn’s satellite
(<3 kbar) and mostly result in minor to moderate
▶ Titan. In addition, it can be present in circum-
changes in the rock’s mineralogy and texture.
stellar environments, for example, in the enve-
(In exceptional cases, however, the original con-
lope of the protoplanetary nebula CRL
stituents are completely replaced.) Commonly,
618 (Cernicharo et al. 2001).
clays, silica minerals, evaporites, or carbonates
replace or alter detrital grains, form cement, and
fill pore spaces, thereby reducing the porosity and
See Also
permeability of the rock.
Diagenesis is an important process controlling
▶ Giant Planets
the preservation of morphological, molecular,
▶ Infrared Space Observatory
chemical, and biological signatures of life in
older rocks.
▶ Titan
Cernicharo J, Heras AM, Tielens AGGM, Pardo JR, ▶ Biomarkers

Herpin F, Guélin M, Waters LBFM (2001) Infrared
space observatory’s discovery of C4H2, C6H2, and
Benzene in CRL 618. Astrophys J 546:L123–L126 ▶ Sedimentary Rock
636 Diamictite/Diamicton
Diamictite/Diamicton Diamino Acid
Daniele L. Pinti Uwe J. Meierhenrich

GEOTOP Research Center for Geochemistry and Institut de Chimie de Nice (ICN), University
Geodynamics, Université du Québec à Montréal, Nice-Sophia Antipolis, Nice, France
Keywords
Synonyms
Amino acids; Chirality; Murchison meteorite;
Mixtite Peptide nucleic acids
Definition Synonyms
Diamictite is a non-sorted or poorly sorted, Diamino carboxylic acid

noncalcareous, terrigenous ▶ sedimentary rock.
Diamictite contains a wide range of particle sizes,
from gravel to sand, dispersed in a poorly or Definition
unstratified clayey matrix. The non-lithified
equivalent of diamictite is diamicton. Diamictite Diamino acids are chemically named diamino
is believed to form primarily in (sub)glacial envi- carboxylic acids. Diamino acids are molecules
ronments as ground and terminal moraines and that contain at least one carboxylic acid group
may also form as debris flows and in volcanic (COOH) and two amine functional groups (NH2).
environments as lahar deposits. Diamino acids belong to the chemical family of
The astrobiological interest in diamictite is amino acids. Diamino acids often but not always
due to its possible significance as climate indica- contain a stereogenic center and can therefore be
tor in suspected glacial sedimentary deposits chiral. The protein ▶ amino acid lysine is a
(▶ Snowball Earth), thus serving to identify sim- diamino acid.
ilar processes in Martian polar caps. The finely
ground continental material in glacially related
diamictites is subject to rapid outwash during Overview
deglaciation and has been invoked to have caused
excess alkalinity and evolution-enhancing nutri- In biochemistry, diamino acids, that is, amino
ent spikes in terminal Cryogenian ecosphere radi- acids containing a second amine functional
ation events. group, are of particular interest. Lysine is a pro-
tein diamino acid. According to systematic stud-
ies on the recruitment order of amino acids and
See Also diamino acids in proteins, diamino acids were
recruited in a relatively late stage of evolution.
▶ Glaciation Ornithine and 2,6-diaminopimelic acid are non-
▶ Polar Caps (Mars) protein diamino acids that play important roles in
▶ Sedimentary Rock the urea cycle and in bacteria cell walls.
▶ Snowball Earth Diamino acids are of particular interest in dis-
▶ Subglacial Environments cussions of chemical evolution and the origin of
Diapirism 637
life on Earth, since they can be used as monomers Muñoz Caro G et al (2002) Amino acids from ultraviolet
for the synthesis of peptide nucleic acids (PNAs), irradiation of interstellar ice analogues. Nature
416:403–406
which some have suggested may have played a Nelson KE et al (2000) Peptide nucleic acids rather than
role in the early evolution of life. There are RNA may have been the first genetic molecule. Proc
numerous possible peptide nucleic acids. One Natl Acad Sci U S A 97:3868–3871
type of PNA molecule is composed of
N-aminoethylglycine leading to aegPNA, and
others can be constructed from diamino acids
(da) leading to daPNA. Peptide nucleic acids Diamino Carboxylic Acid D
such as aegPNA and daPNA are capable of
forming duplex structures with individual RNA ▶ Diamino Acid
and DNA strands. PNAs are therefore nucleic
acid analogs, and they are considered as candi-
dates for the first genetic material on Earth. In this
case, peptide nucleic acids may have transferred Diaminovaleric Acid
the genetic information to RNA and DNA.
The diamino acids 2,3-diaminopropionic acid ▶ Ornithine
and 2,4-diaminobutanoic acid were identified
both in the Murchison meteorite and in the prod-
ucts of comet simulation experiments (artificial
comets). PNA oligomers have never been Diapirism
detected in living organisms, in meteorites, or in
artificial comets, although some research has Nicholas Arndt
suggested these may be prebiotic compounds. ISTerre, Université Grenoble Alpes, France
In the context of the origin of biomolecular
homochirality, the chiroptical properties of
diamino acids have been studied. ▶ Circular Definition
dichroism spectra of zwitterionic L-diamino
acids in solution show maxima at 200 nm. A diapir is a teardrop-shaped body of solid but
ductile, low-density rock that ascends through
denser, more rigid rock. Examples of diapirism
See Also include ice, mud, and salt diapirs in sedimentary
sequences and possibly dome-like intrusions of
▶ Amino Acid granitic gneiss in Archean ▶ cratons. Diapirism
▶ Chirality refers to an upward migration of low-density
▶ Circular Dichroism material, commonly accompanied by downward
▶ Zwitterion migration of denser diapirs nearby. Diapirism
produces a characteristic dome-and-basin struc-
ture that may be characteristic of vertical tecton-
References and Further Reading ics in hot, ductile Archean continental crust. The
Bredehöft JH et al (2007) Chiroptical properties of
higher temperature resulted from the decay of
diamino carboxylic acids. Chirality 19:570–573 radioactive elements, which were more abundant
Meierhenrich U (2008) Amino acids and the asymmetry of in Archean continental crust.
life. Springer, Heidelberg Salt diapirism is widespread on Earth in those
Meinert C et al (2012) N-(2-Aminoethyl)glycine and
amino acids from interstellar ice analogues.
sedimentary basins that had seen the accumula-
ChemPlusChem 77:186–191 tion of thick evaporitic strata, such as Angola,
638 Diastereomers
Morocco, the northern Gulf of Mexico, southern See Also

Iran, western Kazakhstan, and northern Germany.
Mud diapirism is observed in regions where ▶ Amino Acid
overpressure develops in unconsolidated shale- ▶ Chirality
rich strata, such as in pro-delta settings, in ▶ Enantiomers
sabkhas where it is usually related to groundwater ▶ Isomer
fluctuations, and in active hydrocarbon systems.
Mud diapirism may occur on ▶ Mars’s surface
and icy diapirism on Saturn’s satellites.
Diazenylium
See Also William M. Irvine

▶ Archean Tectonics
▶ Craton
▶ Mars
Synonyms
N2H+
Diastereomers
Definition
Jeffrey Bada
Scripps Institution of Oceanography, La Jolla, Protonated molecular nitrogen (protonated
CA, USA ▶ dinitrogen), N2H+, is used by radio astrono-
mers and astrochemists as a tracer of dense gas
in ▶ molecular clouds, since its large
Definition electric dipole moment requires relatively high
densities to collisionally excite the pure rotational
Diastereomers are optically active ▶ isomers that transitions at millimeter wavelengths. Since the
are not ▶ enantiomers. They are characterized by presumed reservoir of most nitrogen in these
having more than one chiral center. clouds, molecular nitrogen (N2), has no easily
A diastereomer with n chiral centers has 2n pos- accessible transitions, N2H+ is an important indi-
sible ▶ stereoisomers. Thus, for a diastereomer cator of the nitrogen chemistry in these regions. It
with two chiral centers, there are four possible is particularly useful, since it is one of the last gas
stereoisomers: a pair of diastereomers with two phase species to freeze out onto the ▶ interstellar
pairs of enantiomers. One example is the diaste- dust grains at the center of cold clouds.
reomers l-isoleucine and d-alloisoleucine. For iso-
leucine, the enantiomers are l- and d-isoleucine
and for alloisoleucine the enantiomers are l- and History
d-alloisoleucine (see Fig. 3 in ▶ Amino Acid).
Diastereomers have different physical proper- The ion N2H+ was identified in interstellar molec-
ties (different melting points, boiling points, sol- ular clouds by means of the characteristic hyper-
ubilities, etc.). Their chemical properties also fine splitting pattern of the fundamental
differ in that they react with reagents at different rotational transition (J = 10) at 93 GHz
rates. Thus, they can be directly separated by (Turner 1974), 2 years before this transition was
chromatographic methods without the need for measured in the laboratory, providing an interest-
chiral reagents. ing example of the ability to do fundamental
Dicarboxylic Acid 639
molecular physics experiments in space. The

interpretation of the emission from this species DIBs
in molecular clouds has been recently reviwed by
Daniel et al. (2013). ▶ Diffuse Interstellar Bands
See Also
Dicarboxylic Acid
▶ Dinitrogen D
▶ Interstellar Dust Henderson James (Jim) Cleaves II
▶ Molecular Cloud Earth–Life Science Institute (ELSI), Tokyo
▶ Molecules in Space Institute of Technology, Meguro–ku, Tokyo, Japan
Washington, DC, USA
Daniel F, Gérin M, Roueff E, Cernicharo J, Marcelino N, Center for Chemical Evolution, Georgia Institute
Lique F, Lis DC, Teyssier D, Biver N, Bockelée- of Technology, Atlanta, GA, USA
Morvan D (2013) Nitrogen isotopic ratios in Barnard 1:
a consistent study of the N2H+, NH3, CN, HCN, and
HNC isotopologues. Astron Astrophys 560:A3
Turner BE (1974) U93.174 – a new interstellar line with Definition
quadrupole hyperfine splitting. Astrophys J 193:
L83–L87 Dicarboxylic acids are organic compounds
containing two ▶ carboxylic acid functional
groups. Dicarboxylic acids generally show the
same chemical behavior and reactivity as mono-
Diazotrophy carboxylic acids. The ionization of the second
carboxyl group occurs less readily than the first
Juli Peretó one, because more energy is required to separate
Institut Cavanilles de Biodiversitat i Biologia a positive hydrogen ion from the anion than from
Evolutiva, Universitat de València, València, the neutral molecule.
Spain Dicarboxylic acids are important metabolic
products, for example, as Krebs cycle intermedi-
ates (e.g., a-ketoglutaric acid, HOOC
Synonyms (CH2)2COCOOH) and the products of fatty acid
oxidation. The simplest dicarboxylic acid is oxalic
Nitrogen fixation acid (HOOCCOOH); others important in bio-
chemistry include malonic (HOOCCH2COOH),
succinic (HOOC(CH2)2COOH), and glutaric
Definition (HOOC(CH2)3COOH) acids. The protein
▶ amino acids, aspartic (HOOCCH2CH(NH2)
Diazotrophy is the metabolic ability to fix atmo- COOH) and glutamic (HOOC(CH2)2CH(NH2)
spheric nitrogen into a biologically useful form COOH) acids, are also dicarboxylic acids.
(i.e., ammonia).
See Also
See Also
▶ Amino Acid
▶ Nitrogen Cycle, Biological ▶ Carboxylic Acid
640 Dichotomy, Planetary
▶ Crust
Dichotomy, Planetary ▶ Mars
▶ Moon, The
Daniela Tirsch
Dicyan
Synonyms ▶ Cyanogen
Separation; Split
Dicyanogen
Definition
▶ Cyanogen
A dichotomy refers to the splitting of surface and
other planetary features in two dissimilar,
nonoverlapping parts. On ▶ Mars, a dichotomy
can be found in terms of crater density and topog-
raphy with the highly cratered southern highlands Diderot’s Conception of Origins
and the less cratered northern lowlands. A further of Life
dichotomy is speculated to exist with respect to
crustal thickness. The ▶ crust of the northern Stéphane Tirard
lowlands could be much thinner (ca. 32 km) Centre François Viète d’Histoire des Sciences et
than the crust of the southern highlands des Techniques EA 1161, Faculté des Sciences et
(ca. 58 km). This has been derived from an inter- des Techniques de Nantes, Nantes, France
pretation of the gravity field assuming a constant
crust density. The two competing formation
hypotheses involve an exogenic cause (e.g., a History
giant impact) and an endogenic formation mech-
anism (e.g., degree-1 mantle convection). On the The French philosopher Denis Diderot
Moon, a dichotomy is known regarding the dis- (1713–1884) wrote about life in several of his
tribution of ▶ basalt, with the near side compris- more famous texts, notably in his Entretiens
ing about 90 % whereas the far side only exhibits (1776) and especially in Le Rêve de d’Allembert
10 % of the bulk basalt abundance. The reason for (from 1769, but published in 1830). Diderot was a
this is again a dichotomy in crustal thickness partisan of spontaneous generation and of trans-
causing basalt plumes to easily reach the surface formation of matter theories. In his Elements de
on the thinner near side (60 km) in contrast to the Physiologie (published posthumously in 1875),
thicker far side (120 km). The Earth’s hypsomet- he claimed that nature produced only a few living
ric curve displays another dichotomy in surface beings, perhaps only one, of which all the others
elevation between the continents and the ocean would come from by combination, micturition,
basins. and dissolution and which would vary infinitely.
See Also See Also
▶ Basalt ▶ Buffon’s Conception of Origins of Life

▶ Crater, Impact ▶ Lamarck’s Conception of Origins of Life
Differentiation, Planetary 641
are the release of energy through gravitational

Differentiation separation of light and dense constituents, the
decay of radioactive heat sources, and additional
▶ Fractionation external heat sources such as tidal heating (e.g.,
Schubert et al. 1986; see ▶ Tides, Archean). Pro-
vided that a sufficient amount of energy was
retained during accretion, a near-surface magma
Differentiation, Planetary ocean would have formed atop of the growing
protoplanet. Chemical differentiation of the D
1 2
Frank Sohl and Doris Breuer magma ocean would be affected by impact stir-
1
Deutsches Zentrum f€ur Luft- und Raumfahrt ring, convective mixing, cooling, and solidifica-
ur Planetenforschung, Berlin, tion (e.g., Abe 1993). The solidification timescale
Germany of the magma ocean is determined by the balance
2
German Aerospace Center (DLR), Institute of between the heat flow entering from below, latent
Planetary Research, Berlin, Germany heat release upon crystallization, and heat loss at
the surface. The crystallization of the magma
ocean is thought to cause initial compositional
Keywords layering in terms of unstable cumulate density
stratification, thereby invoking gravitational
Accretion; Convective overturn; Core formation; overturn by convective instability (Elkins-Tanton
Crust segregation; Degassing; Magma ocean; et al. 2003). This may have led to the formation of
Partial melt an early basaltic crust and chemically distinct
mantle reservoirs on terrestrial planets. Aqueous
alteration involving chemical reactions between
Definition olivine-rich mantle ▶ rock and liquid water may
have caused the release of a substantial amount of
Planetary differentiation is the separation of dif- ▶ hydrogen (▶ Serpentinization). If the energy
ferent constituents of planetary materials released by gravitational separation of light and
resulting in the formation of distinct composi- dense constituents were sufficient to invoke melt-
tional layers. Denser material tends to sink into ing of the remaining mantle material, the separa-
the center and less dense material rises toward the tion process would become energetically self-
surface. If internal differentiation of a ▶ terres- sustaining and will run to completion within a
trial planet has run to completion, the interior short period of time (tens of million years). The
would be subdivided into an iron-rich metallic subdivision of terrestrial planet interiors into
core and a silicate ▶ mantle, overlain by a chem- crust, mantle, and core (e.g., Sohl and Schubert
ically distinct ▶ crust enriched in ▶ basalt. 2007; see ▶ Interior Structure, Planetary) can be
attributed to such intense differentiation events,
commonly referred to as runaway differentiation.
Overview
The accretion of the terrestrial bodies from plan- Basic Methodology

etesimals provides the initial source of energy for
subsequent planetary evolution. The amount of The differentiation and associated chemical evo-
kinetic energy converted to heat during accretion lution of a terrestrial body is in general the con-
is not well known because it is strongly depen- sequence of partial melting in its interior. Density
dent on specific formation timescales and envi- differences between the melt and the solid resid-
ronmental conditions. Heat sources, other than uum would then result in their separation. For
deposition of impact energy during accretion, larger planets with increasing accretion time,
642 Differentiation, Planetary
the melting is caused by impact energy and ancient asteroids that have been shattered by
results in the formation of a magma ocean close impacts. Studies on short-lived radionuclides
to the surface. In a magma ocean, the much (e.g., 182Hf) suggest that the core formation pro-
denser iron sinks and accumulates at its base. cess occurred during the first 5–30 million years
Iron parcels then settle further to the center of of the solar system, prolonging with increasing
the solid or partly solid planet by Rayleigh- size of the planetary body (e.g., Kleine
Taylor instability (Stevenson 1990). The larger et al. 2002). Thus, the differentiation event
the accumulated iron diapirs, the higher are the occurred most likely contemporaneously or
sinking velocities. The gravitational energy shortly after completion of ▶ asteroid or planet
released by core formation is converted into ther- accretion. The relatively rapid separation of sili-
mal energy, which strongly heats the interior. As cate and iron implies that both components were
a consequence, the temperature profile is inverted fluid (Stevenson 1990). In case of small planetary
to decrease from the center toward the surface. bodies like the asteroids, the decay of short-lived
The total energy released by the differentiation of radioactive isotopes like 26Al and 60Fe provides a
a homogeneous planet into an iron-rich core and a sufficient amount of energy to melt the interior
silicate mantle can be estimated from the differ- (e.g., MacPherson et al. 1995; Tachibana and
ence between the potential energy stored in a Huss 2003). The presence of siderophile ele-
homogeneous planet after accretion and the ments, i.e., elements such as S, Ni, and Co that
potential energy of the differentiated, two-layer are readily soluble in molten iron, would result in
planet (Schubert et al. 1986). If that amount of a significant melting-point depression (e.g.,
energy were distributed homogeneously within Braginsky 1964), thereby facilitating the segre-
the interior, assuming thermal equilibrium gation of iron alloys in the core. Once a metal
between the mantle and the core, the mean tem- core has formed, it may further differentiate as
perature would rise about 1,000 K for the Earth soon as an inner pure Fe core starts to solidify
and about 300 K for Mars (Solomon 1979). upon cooling. In turn, the fluid outer core then
Further chemical differentiation of a planetary would be enriched in the low-melting-point
mantle is usually associated with crust formation. components.
The process of crust formation can, in general, be The formation of the secondary crust would
divided into two phases: primary and secondary deplete the mantle in radioactive elements and
crust formation. A primary crust can be formed as crust-forming minerals, as those are incompatible
a consequence of freezing of a magma ocean and enriched in the melt. In addition, the melt that
during an early state of planetary evolution. forms the crust is enriched in volatiles that are
Whereas light minerals like plagioclase feldspar released into the atmosphere by extrusive volca-
would rise toward the surface forming an anor- nism. Mantle degassing associated with crustal
thositic crust, heavy minerals such as olivine differentiation can play an important role for the
would sink to the bottom of the magma ocean. evolution of planetary atmospheres. The mecha-
This gravitationally induced crystal differentia- nisms of crust formation differ between ▶ plate
tion will result in a chemically layered mantle. tectonics and one-plate planets. Plate tectonics
provides for crust formation in a two-step fash-
ion. Crust formation is most effective at divergent
Key Research Findings plate boundaries, where rising hot mantle mate-
rial crosses the solidus near the surface. The
The separation of the metal from the silicate corresponding pressure-release melting generates
particles is a common process in the ▶ solar basaltic crust that is continuously recycled at
system. Prominent examples are the iron-rich convergent plate boundaries. More silicic crust
cores of the terrestrial bodies and iron ▶ meteor- is produced in a second differentiation step at
ites that consist of Fe-Ni alloys. The latter are convergent plate boundaries where basaltic crust
assumed to be the fragments of the cores of larger is remelted (together with wet continental
Differentiation, Planetary 643
sediments and possibly mantle rock) to form new interior of main-belt asteroid 4 Vesta (▶ Vesta) is
continental-like crust (e.g., Taylor and likely differentiated (Ghosh and McSween 1998),
McLennan 1985). One important open question as suggested by the spectrally variegated surface
for the Earth with its plate tectonics is the timing of that accretional remnant.
of the continental crust growth. Models range
from rapid early crustal growth with crust pro-
duction and recycling in equilibrium in the later Future Directions
evolution (Armstrong 1981), a continuous
growth of the crust to its present size (Veizer To further our understanding of the internal dif- D
and Jansen 1979), and an episodic growth with a ferentiation and the associated volatile loss, new
peak at the Archaean-Proterozoic transition and improved techniques and methods will be
(Taylor and McLennan 1985). For one-plate developed and applied in the future. Among
planets in the ▶ stagnant lid convection regime, those are isotopic and trace element studies on
crust recycling is only possible in the form of planetary material, improved numerical tech-
crustal delamination for which the lower-dense niques incorporating mantle melting and
crustal material (basalt that changes to eclogite at degassing in the frame of three-dimensional con-
depth) sinks into the mantle and there is no vection models, and also future missions apply-
two-stage differentiation. Instead, melt is formed ing new and improved measuring techniques.
underneath the lithosphere usually at greater
depth as compared to planets with plate tectonics.
The crust produced would predominately be See Also
composed of basaltic rock. Observations and
thermochemical models show that for the Moon ▶ Anorthosite
and ▶ Mars, most of the basaltic crust is produced ▶ Archean Eon
early in the planetary evolution (e.g., Nimmo and ▶ Asteroid
Tanaka 2005). ▶ Venus is distinct in that the ▶ Asteroid Belt, Main
surface has an average age of 300–800 Ma ▶ Basalt
(Schaber et al. 1992). The uniform age distribu- ▶ Core, Planetary
tion has been associated with a global resurfacing ▶ Crust
event, possibly associated with a change in the ▶ Diapirism
style of mantle convection (see Bougher ▶ Heat Flow, Planetary
et al. (1997) for a review). The relatively rapid ▶ Hydrogen
global resurfacing of Venus may be questioned, ▶ Interior Structure, Planetary
since the cratering record allows a variety of ▶ Mantle
interpretations in terms of volcanic resurfacing, ▶ Mars
including a global decrease in activity over time ▶ Meteorites
(Campbell et al. 1999). ▶ Moon, The
▶ Planetesimals
▶ Plate Tectonics
Applications ▶ Primordial Heat
▶ Radioactive Heating
The state of knowledge of planetary differentia- ▶ Rock
tion processes has important implications for the ▶ Satellite or Moon
general understanding of the internal structure, ▶ Serpentinization
thermochemical evolution, bulk composition, ▶ Solidus
and atmosphere evolution of the terrestrial ▶ Stagnant Lid Convection
planets and terrestrial-type bodies like the Moon ▶ System Solar Formation, Chronology of
and several large outer-planet satellites. Even the ▶ Terrestrial Planet
644 Diffraction
▶ Tides, Planetary Sohl F, Schubert G (2007) Interior structure, composition, and

▶ Venus mineralogy of the terrestrial planets. In: Schubert G (ed-in-
chief) Treatise on geophysics. Spohn T (vol ed) Planets
▶ Vesta and moons, vol 10. Elsevier, Amsterdam, pp 27–68
Solomon SC (1979) Formation, history and energetics of
cores in the terrestrial planets. Phys Earth Planet Inter
19:168–182
References and Further Reading Stevenson DJ (1990) Fluid dynamics of core formation.
In: Newsom HE, Jones JH (eds) Origin of the Earth.
Abe Y (1993) Thermal evolution and chemical differenti- Oxford University Press, New York, pp 231–249
ation of the terrestrial magma ocean. In: Takahashi E, Tachibana S, Huss GI (2003) The initial abundance of
Jeanloz R, Rubie D (eds) Evolution of the earth and 60Fe in the solar system. Astrophys J 588:L41–L44
planets, geophys, vol 14, Monograph Series 74 and Taylor SR, McLennan SM (1985) The continental crust:
IUGG. American Geophysical Union and Interna- its composition and evolution. Blackwell Scientific,
tional Union of Geodesy and Geophysics, Washington, Oxford
DC, pp 41–54 Veizer J, Jansen SL (1979) Basement and sedimentary
Armstrong RL (1981) Radiogenic isotopes. The case for recycling and continental evolution. J Geol 87:341–370
crustal recycling on a near-steady-state no-
continental-growth Earth. Roy Soc Lond
Philipp Trans A301:433–471
Bougher SW, Hunten DM, Phillips RJ (eds) (1997) Space
science series: Venus II – geology, geophysics, atmo-
sphere, and solar wind environment. The University of
Diffraction
Arizona Press, Tucson, p 1330
Braginsky SI (1964) Magnetohydrodynamics of the Earth Daniel Rouan
core. Geomagn Aeron 4:698–712 LESIA, Observatoire Paris-Site de Meudon,
Campbell BA, Campbell DB, DeVries CH (1999) Surface
Meudon, France
processes in the Venus highlands: results from analysis
of Magellan and Arecibo data. J Geophys Res
104:1897–1916
Elkins-Tanton LT, Parmentier EM, Hess PC Definition
(2003) Magma ocean fractional crystallization and
cumulate overturn in terrestrial planets: implications
for Mars, meteoritics and planet. Science 38: Diffraction, in physics, is a phenomenon that
1753–1771 occurs when a wave encounters an obstacle or
Ghosh A, McSween HJ Jr (1998) A thermal model for the travels through a medium with a varying refractive
differentiation of asteroid 4 Vesta, based on radiogenic
index. Secondary radial waves issued from the
heating. Icarus 134:187–206
Joliff BL, Wieczorek MA, Shearer CK, Neal CR obstacle spread out, the more widely as the size
(2006) New views of the Moon. Mineralogical Society of the obstacle becomes smaller and approaches
of America, Reviews in Mineralogy and Geochemis- the wavelength. Because the secondary waves
try, vol 60, Washington, DC
may interfere, the diffraction pattern generally
Kleine T, M€unker C, Mezger K, Palme H (2002) Rapid
accretion and early core formation on asteroids and the shows structures with fringes or rings (see
terrestrial planets from Hf-W chronometry. Nature Fig. 1). Diffraction of light in observational astron-
418:952–955 omy limits the ability of an imaging system to
MacPherson GJ, Davis AM, Zinner EK (1995) The distri-
bution of aluminium-26 in the early solar system – a
resolve the finest details. The larger the aperture
reappraisal. Meteoritics 30:365–386 of the telescope and the smaller the wavelength,
Nimmo F, Tanaka K (2005) Early crustal evolution of the finer the angular resolution. A quantitative
Mars. Annu Rev Earth Planet Sci 33:133–161 expression of this limit is given by the expression
Schaber GG, Strom RG, Moore HJ et al (1992)
y(arcsec) = 0.2 l(mm)/D(m), where l is the
Geology and distribution of impact craters on Venus:
what are they telling us? J Geophys Res wavelength in mm and D the diameter of the tele-
97:13,257–13,301 scope in meters. This effect provides one justifica-
Schubert G, Spohn T, Reynolds RT (1986) Thermal his- tion for telescopes having very large mirrors, the
tories, compositions and internal structures of the
other justification being better collecting power.
moons of the solar system. In: Burns JA, Shapley MS
(eds) Satellites. University Arizona Press, Tucson, However, on the ground (as opposed to space),
pp 224–292 telescopes are hampered by atmospheric
Diffuse Cloud 645
Diffraction,
Fig. 1 Diffraction pattern
generated by a small hole in
a dark screen illuminated
by a beam of light and seen
at a certain distance. Note
the rings that are
characteristic of this figure
called the Airy pattern
D
turbulence that blurs the image to a typical resolu- 1 magnitude, corresponding typically to densities
tion of 1 arcsec, whatever the size of the telescope of a few tens of atoms per cubic centimeter or
is. Only ▶ adaptive optics techniques allow to less, and column densities of order 1021 hydrogen
recover the diffraction limit of a telescope. atoms cm2. Hydrogen, the principal constituent
of interstellar clouds, is predominantly in atomic
form in diffuse clouds. The physics and chemis-
See Also try of diffuse clouds may be studied by observing
the absorption lines produced in the spectra of
▶ Adaptive Optics background stars. Trace constituents observed at
▶ Coronagraphy visual and ultraviolet wavelengths include many
▶ Imaging atoms and diatomic molecules, including such
simple molecules as CO, OH, CH, CH+, CN,
and H2. At millimeter wavelengths, molecules
such as C2H and HCO+ are observed. As the
density increases, diffuse clouds grade gradually
Diffuse Bands
into those clouds referred to as translucent and
then to those that are called dense. The Herschel
▶ Diffuse Interstellar Bands
satellite has recently discovered OH+, H2O+,
H2Cl+, and HF as constituents of diffuse inter-
stellar clouds (Neufeld et al. 2010; Gerin
et al. 2010; Lis et al. 2010).
Diffuse Cloud
William M. Irvine
University of Massachusetts, Amherst, MA, USA See Also
▶ Dense Cloud
Definition ▶ Herschel Mission
A diffuse cloud is that portion of an interstellar ▶ Interstellar Medium
cloud with visual extinction (Av) less than about ▶ Molecules in Space
646 Diffuse Galactic Light

Diffuse Interstellar Bands
Gerin et al (2010) Interstellar OH+, H2O+ and H3O+ along
the sight-line to G10.6-0.4. Astron Astrophys 518:
Martin A. Cordiner
L110–L114
Lis D et al (2010) Herschel-HIFI discovery of The Goddard Center for Astrobiology, NASA
interstellar chloronium (H2Cl+). Astron Astrophys Goddard Space Flight Center, Greenbelt,
521:L9–L113 MD, USA
Neufeld D et al (2010) Strong absorption by
interstellar hydrogen fluoride: Herschel-HIFI observa-
tions of the sight-line to G10.6-0.4 (W31C). Astron
Astrophys 518:L108–L112 Keywords
Snow TP, McCall BJ (2006) Diffuse atomic and
molecular clouds. Ann Rev Astron Astrophys 44:
Astrochemistry; Interstellar dust; Interstellar
367–414
medium; Large molecules; Optical spectroscopy
Synonyms
Diffuse Galactic Light DIBs; Diffuse bands
William M. Irvine
University of Massachusetts, Amherst, MA, USA Definition
The diffuse interstellar bands (DIBs) are broad

spectroscopic absorption features of unknown
origin, which occur in the optical and near-
Definition
infrared spectra of stars as their light passes
through the interstellar medium. More than
The diffuse galactic light is the weak
400 DIBs are currently known, at least some of
background ultraviolet and visible radiation
which are believed to be caused by large carbon-
produced by ▶ interstellar dust grains
based molecules such as polycyclic aromatics,
scattering light from stars in the disk of the
carbon chains, and ▶ fullerenes.
▶ Milky Way galaxy. It is not easily
separable observationally from the ▶ zodiacal
light (sunlight scattered by dust in the solar
History
system) and from airglow (emission from
atoms and molecules in the Earth’s upper
In 1922, Mary Lea Heger discovered two prom-
atmosphere).
inent dark absorption bands in the spectra of
early-type stars at wavelengths of 5,780 and
5,797 Ångstrom (Heger 1922). The cause of
these features could not be established.
See Also Due to the detailed studies of binary stars by
Merrill and Sanford (1938), the unidentified
▶ Background bands were determined to be of interstellar origin,
▶ Interstellar Dust and four additional bands were discovered in the
▶ Zodiacal Light red region of the spectrum, at wavelengths of
Diffuse Interstellar Bands 647
6,203, 6,270, 6,284, and 6,614 Å. The bands were chemical conditions in order to elucidate the
noted as being somewhat diffuse, as they have a nature of the carriers. The spectra are carefully
broader width than other interstellar absorption compared with laboratory and theoretical data to
lines that are known to be caused by gas-phase test hypotheses regarding potential carriers. To
atoms (such as sodium and calcium) or simple assign a DIB carrier is very difficult because it
molecules (such as CH and CN). Henceforth, requires a precise spectroscopic match between
these and other unidentified, broad interstellar the DIB and the proposed carrier spectrum, at
absorption features in optical spectra came to be temperature, density, and radiation conditions
known as diffuse interstellar bands (DIBs). matching those of the diffuse ▶ interstellar D
More than 400 different DIBs are now known, medium (ISM) where the carriers reside.
with wavelengths 400–1,800 nm, and more are
found with each increase in survey sensitivity.
The carriers of these spectral features are yet to Key Research Findings
be definitively assigned.
Observational Constraints on the DIB Carriers
Throughout our Galaxy, the strengths of many of
Overview the diffuse interstellar bands are closely corre-
lated with the amount of ▶ interstellar dust,
The diffuse interstellar band problem is among the which causes reddening of starlight. This sug-
longest-standing unsolved mysteries in astrophys- gests a possible origin in small solid particles
ics and has confounded the efforts of astronomers that are known in the laboratory to exhibit broad
and chemists since the early twentieth century. At optical absorption lines. Diffuse interstellar
least some of the DIBs are probably attributable to bands are observed in low-density regions of the
large, carbon-based molecules. The resolution of interstellar medium whenever dust is present and
this mystery is therefore likely to be of relevance to even in regions pervaded by intense, hard ultra-
theories regarding the evolution of organic mole- violet radiation that rapidly destroys all but the
cules in space and may therefore be of relevance hardiest molecular material. However, the corre-
for astrobiology. Due to their ubiquity in the dusty lation between DIB strengths and amount of dust
interstellar medium and their chemical stability in is imperfect, with a scatter of about a factor of
harsh environments, the DIB carriers constitute a 2 about the mean for a given reddening for the
possible source of cosmic carbon delivered to the well-studied 5,780 Å DIB (Herbig 1993), which
early Earth. constitutes evidence against a solid-state carrier
particle. In addition, the relative constancy of the
DIB wavelengths across different astrophysical
Basic Methodology environments, the lack of any light polarization
signature inside the DIB profiles (the interstellar
Diffuse interstellar bands are most readily dust is known to produce polarization), and the
detected and analyzed in high-resolution, high lack of emission adjacent to the DIBs make inter-
signal-to-noise optical spectra of hot stars (with stellar dust an unlikely carrier. Discussions on the
spectral types O and B). The interstellar bands viability of various carrier candidates are given
may be distinguished from stellar lines through by Herbig (1995), Snow (1995), and Sarre (2006).
multi-epoch observations of spectroscopic binary Strong evidence that several DIBs are caused
stars or by comparison with un-reddened stan- by gas-phase molecules is given by the observa-
dard stars. Variations in DIB spectra may be tion of diffuse bands in emission in the Red
studied with respect to interstellar physical and Rectangle nebula (Sarre 1991; Fossey 1991).
648 Diffuse Interstellar Bands
The wavelengths of these emission bands are best-studied DIBs are found at rest wavelengths
close to those of a subset of relatively narrow of 5,780 and 5,797 Å. Spectra of these DIBs are
diffuse interstellar bands, and the steep blue shown in Fig. 1, observed toward stars in
(short wavelength) side and red-degraded tails the Milky Way, the Andromeda Galaxy, and the
are typical of electronic transitions in large, cold Large Magellanic Cloud (LMC).
molecules.
Further evidence for large, carbonaceous car- DIB Carrier Candidates
rier molecules is given by close examination of Carbon is unique in its ability to form large,
the shapes of many of the strongest diffuse inter- complex molecules. It is also one of the most
stellar bands, which reveal complex profiles with abundant chemical species in the interstellar
fine structure similar to that seen for various medium, where it exists in the form of neutral
classes of large (gas-phase) molecules in the lab- and ionized atoms, polyatomic molecules, aro-
oratory, including long carbon chains, ▶ polycy- matics, and dust grains. The fact that there are
clic aromatic hydrocarbons (PAHs), and so many different DIBs, each apparently caused
▶ fullerenes. The fine structure of these DIB pro- by a different, large gas-phase molecule, is
files is observed to be variable, perhaps as a result strongly suggestive that the molecular structures
of changes in the internal temperature, excitation, of at least some and perhaps the majority of the
or isotopic composition of the carriers. DIB carriers are composed predominantly of carbon.
widths are consistent with carrier molecules Large, carbon-based molecules, such as
containing at least about 20 atoms. ▶ polycyclic aromatic hydrocarbons (and the
DIB strengths correlate well with the amount various classes of PAHs that have been modified,
of neutral gas (such as atomic hydrogen, H I, and for example, by ionization, protonation, or dehy-
Na I), but not with the denser molecular gas drogenation), are stable and resistant to destruc-
traced by H2, as demonstrated by Herbig (1993). tion by UV radiation. Such aromatic molecules
The DIB carriers are not, therefore, closely asso- are believed to be abundant in the diffuse ISM
ciated with the chemistry of those molecules that where DIBs are strongest and therefore represent
are typically most abundant in dense molecular an attractive class of possible carriers. This
environments where stars form and UV radiation hypothesis has been reviewed by several authors
fields are low. Indeed, various studies have shown (Crawford et al. 1985; Leger and D’Hendecourt
that DIBs tend to be relatively weak inside dense 1985; Salama et al. 1999). Laboratory spectros-
molecular clouds (e.g., Adamson et al. 1991). copy has shown that PAHs and their ions possess
The fact that DIBs are observed most strongly strong electronic transitions in the optical (e.g.,
in the relatively unshielded ▶ diffuse clouds with Ruiterkamp et al. 2002; Tan and Salama 2006;
low extinction indicates that the carriers are resis- Useli-Bacchitta et al. 2010). However, after
tant to destruction by UV radiation, so they can- extensive laboratory and theoretical studies of
not be small molecules, which tend to be easily the spectroscopy of a plenitude of PAHs and
photodissociated. related aromatics, no DIB has yet been assigned
Research into the properties of diffuse inter- to any specific PAH-type molecule.
stellar bands outside the Milky Way (MW) is Following the suggestion of Kroto in the
currently in progress (see, e.g., Cox et al. 2006; 1980s that C60 and other ▶ fullerenes might be
Cordiner et al. 2011a), and although the DIB present in the ISM (see Kroto and Jura 1992),
strengths show some deviation from the trends Ehrenfreund and Foing (1997) examined the pos-
observed in the Milky Way (presumably as a sibility of C 60 + as the carrier of two DIBs in the
result of the differing physical and chemical con- near infrared. Other related large molecules that
ditions in these environments, including the have transitions in the optical include carbon
▶ metallicity and interstellar radiation field nanotubes, “Bucky onions,” and graphene sheets.
strength), their overall spectrum is similar Such stable, carbon-based molecules, which span
to that observed in our Galaxy. Two of the the continuum of sizes from large molecules to
Diffuse Interstellar Bands 649
Diffuse Interstellar Bands, Fig. 1 The 5,780 and et al. (2006)). Spectra have been normalized and offset
5,797 Å diffuse interstellar bands (DIBs) observed toward vertically for display. The MAG 63885 data have a fitted
the Milky Way (MW) star b1 Sco (top), the Andromeda profile overlaid in red. The M31 and LMC DIBs are
(M31) star MAG 63885 (middle), and the Large shifted due to the Doppler motion of those galaxies, as
Magellanic Cloud (LMC) star Sk 69 223 (bottom) indicated by the horizontal velocity scale at the top of the
(Data are from Cordiner et al. (2008) and Cox graph
small dust grains, may be injected into interstellar C2H2 and HCN in stellar outflows (Hudgins
space in the outflows from dying stars, later to be et al. 2005). Although they are less stable to UV
incorporated into the next generation of star- and degradation than PAHs, larger nitrogen heterocy-
planet-forming regions. cles (containing at least 50 carbon atoms) should
Theoretically, the addition or substitution of be able to survive in the diffuse ISM to act as
other atoms inside or at the periphery of aromatic possible DIB carriers.
carbon structures, either during the formation of Heteroatoms can also be added to large carbo-
the molecules or during their subsequent journey naceous molecules in unusual ways to modify
through the interstellar medium, results in mole- their optical properties. For example, iron atoms
cules of biological relevance that may constitute bond to the delocalized electron clouds above
potential DIB carriers. For example, the alkyl- aromatic rings to form the so-called organome-
ation and oxidation of aromatic molecules in tallics. During the formation of fullerenes, metal
interstellar ice analogs (Elsila et al. 2006) results atoms have a propensity to become trapped inside
in the addition of methyl and hydroxyl side the carbon cage. Such species highlight the diver-
groups of the kind found in biomolecules. Het- sity of molecules that must be considered in the
erocyclic aromatic organic compounds, including search for plausible DIB carrier candidates.
biologically important nitrogen heterocycles, are Some DIBs may be due to smaller organic
probably also present in the ISM and are theo- molecules; close spectroscopic matches of the
rized to be produced from reactions involving 8,037 Å DIB with CH2CN and the 4,881 and
650 Diffuse Interstellar Bands
5,450 Å DIBs with l-C3H2 have been identified Cordiner MA, Smith KT, Cox NLJ, Evans CJ, Hunter I,
by Cordiner and Sarre (2007) and Maier Przybilla, N, Bresolin F, Sarre PJ (2008)
Diffuse Interstellar Mands in M33. Ast Astrophys
et al. (2011), respectively. Further confirmation 492:L5–L8
is required, however, before these can be consid- Cordiner MA, Cox NLJ, Evans CJ, Trundle C, Smith KT,
ered to be carrier assignments. Sarre PJ (2011a) Astrophys J 726:39–60
Cordiner MA, Cox NLJ, Trundle C, Evans CJ, Hunter I,
Przybilla N, Bresolin F, Salama F (2011b) Astron
Astrophys 480:L13–L16
Applications Cox NLJ, Cordiner MA, Cami J, Foing BH, Sarre PJ,
Kaper L, Ehrenfreund P (2006) Astron Astrophys
Diffuse interstellar bands are clearly visible in 447:991–1009
Crawford MK, Tielens AGGM, Allamandola LJ
reddened spectra of luminous objects throughout (1985) Astrophys J 239:L45–L48
the universe. Once identified, the ubiquity of Ehrenfreund P, Foing BH (1997) Adv Space Res
the DIBs will probably make them invaluable 19:1033–1042
as probes of interstellar chemistry and physics. Elsila JE, Hammond MR, Bernstein MP, Sandford SA,
Zare RN (2006) Meteoritics 41:785–796
Presently, their strengths are useful measures Fossey SJ (1991) Nature 353:6343–6393
of the amount of interstellar dust in a Heger M (1922) Lick Obs Bull 10:146–147
sightline and as tracers of large organic mole- Herbig GH (1993) Astrophys J 407:142–156
cules in space. Herbig GH (1995) Annu Rev Astron Astrophys
33:19–74
Hobbs LM, York DG, Thorburn JA, Snow TP, Bishof M,
Friedman SD, McCall BJ, Oka T, Rachford B,
Future Directions Sonnentrucker P, Welty DE (2009) Astrophys
J 705:32–45
Hudgins DM, Bauschlicher CW Jr, Allamandola LJ
Further studies are required on how variations in (2005) Astrophys J 632:316–332
interstellar physical and chemical conditions Kroto HW, Jura M (1992) Astron Astrophys 263:275–280
affect the DIB spectrum in order to further eluci- Leger A, D’Hendecourt L (1985) Astron Astrophys
date the properties of the carriers. In particular, 146:81–85
Maier JP, Walker GAH, Bohlender DA, Mazzotti FJ,
the examination of peculiar or extreme interstel- Raghunandan R, Fulara J, Garkusha I, Nagy A (2011)
lar environments in our Galaxy and beyond is Astrophys J 726:41–50
likely to yield information in this regard. Con- Merrill PW, Sanford RF (1938) Astrophys
tinuing dedicated laboratory and theoretical J 87:118–132
Ruiterkamp R, Halasinski T, Salama F, Foing BH,
efforts are necessary to determine the optical Allamandola LJ, Schmidt W, Ehrenfreund P (2002)
spectra of candidate carriers in order to make or Astron Astrophys 390:1153–1170
rule out DIB assignments. Salama F, Galazutdinov GA, Krełowski J,
Allamandola LJ, Musaev FA (1999) Astrophys
J 526:265–273
Sarre PJ (1991) Nature 351:6325–6356
See Also Sarre PJ (2006) J Mol Spectrosc 238:1–10
Snow T (1995) A critique of suggested diffuse band
carriers. In: Tielens AGGM, Snow TP (eds) The dif-
▶ Diffuse Cloud fuse interstellar bands, Astrophysics and space
▶ Fullerene science library, vol 202. Kluwer, Dordrecht,
▶ Interstellar Dust pp 379–393
▶ Polycyclic Aromatic Hydrocarbon Tan X, Salama F (2006) Cavity ring-down spectroscopy of
jet-cooled 1-pyrenecarboxyaldehyde (C17H10O) and
1-methylpyrene (C17H12) cations. Chem Phys Lett
422:518–525
References and Further Reading Tielens AGGM, Snow TP (eds) (1995) The diffuse inter-
stellar bands, Astrophysics and space science library,
Adamson AJ, Whittet DCB, Duley WW (1991) Mon Not vol 202. Kluwer, Dordrecht
R Astron Soc 252:234–245 Useli-Bacchitta F, Bonnamy A, Mulas G, Malloci G,
Cordiner MA, Sarre PJ (2007) Astron Astrophys Toublanc D, Joblin C (2010) Chem Phys 371:16–23
472:537–545
Diketopiperazine 651
Definition
Diffusion
Dihydroxyacetone (IUPAC name
Sylvia Ekström 1,3-Dihydroxy-2-propanone) is one of the three
Observatoire Astronomique de l’Université de simplest monosaccharides (“sugars”). Like the
Genève, Faculté des Sciences, Université de two others (L- and D-glyceraldehyde), it
Genève, Versoix, Switzerland has three carbon atoms and is called a
triose. But the molecule is symmetric and
hence has no chiral center. It is present in cells D
Definition where it plays a role in ATP production and
other biological pathways and is produced by
Diffusion refers to an irreversible transport process ▶ glycolysis. Dihydroxyacetone has been sought
acting at the molecular scale and by which the density in the interstellar medium, but there is no con-
of a quantity X (e.g., the mass fraction of a chemical vincing detection up to now.
element) changes with time. The driving mechanism
for the mixing is generally the density gradient ∇X,
and the result is to homogenize the density through-
out the considered region. The general form of the See Also
equation of diffusion can be written as
▶ Chirality
@ ▶ Glycolaldehyde
ðrXÞ ¼ ∇ ðrD∇XÞ
@t ▶ Molecules in Space
with r the mass density of the medium (with units
[mass length3]) and D the diffusion coefficient
(with units [length2 time1]) or diffusivity. The
characteristic timescale for diffusion is: Diketopiperazine
‘
2
tdiff / Koichiro Matsuno
D
Nagaoka University of Technology, Nagaoka,
with ‘ the characteristic size of the considered Japan
region.
Keywords
Diffusion Potential
Amino acids; Anhydride; Cyclic dimer;
▶ Membrane Potential Oligopeptide
Dihydroxyacetone Synonyms
Didier Despois Glycine anhydride
CNRS-Universite de Bordeaux, France
Definition
Synonyms
A diketopiperazine is a cyclic dimer of two
DHA; HOCH2COCH2OH peptide-bonded ▶ amino acid molecules.
652 Dimethyl Ether
Overview which both monomeric amino acids and diketopi-

perazines are supplied in abundance, a wide vari-
Amino acids, delivered via extraterrestrial deliv- ety of oligopeptides could be synthesized. Some
ery or synthesized by Miller-Urey-type chemis- functional oligopeptides could be formed, even at
try, may have been available in abundance on the the stage prior to the emergence of nucleotide or
primitive Earth and may be abundant via these oligonucleotide molecules serving as reaction
sources on other planets as well. Subsequent templates.
chemical evolution would require the synthesis
of oligopeptides from the amino acid monomers.
If a source of energy driving the formation of
peptides is available in the form of geothermal See Also
heat exposed to the ocean as in hydrothermal
environments on Earth, diketopiperazines could ▶ Amino Acid
quite easily be synthesized with the aid of the heat ▶ Hydrothermal Reaction
energy. The ease with which diketopiperazine is ▶ Oligopeptide
formed is a result of the presence of both a free ▶ Origin of Life
amino group and a free carboxylic group residing
in a single dipeptide once the latter is synthesized.
A dipeptide having only one peptide bond linking
its two units initially is unstable against transfor- Nagayama M, Takaoka O, Inomata K, Yamagata Y (1990)
mation into a diketopiperazine with two peptide Diketopiperazine-mediated peptide formation
bonds in a closed-ring form. in aqueous solution. Orig Life Evol Biosph
20:249–257
At first sight, diketopiperazine may look like a
dead end for chemical evolution since there is no
room for making additional peptide bonds.
However, this is a hasty conclusion. There is
certainly a possibility for opening up the
once-closed ring of peptide bonds when Dimethyl Ether
the surrounding conditions are changed. If the
amount of diketopiperazine synthesized in the William M. Irvine
reaction solution becomes sufficiently high, University of Massachusetts, Amherst, MA, USA
the reaction of an amino acid molecule
aminolyzing a diketopiperazine to form a
tripeptide is actually confirmed experimentally Synonyms
(Nagayama et al. 1990). The ring opening is due
to the nucleophilic attack of an amino acid mol- CH3OCH3; Methoxymethane (IUPAC name)
ecule’s amino group on the diketopiperazine. The
similar nucleophilic attack on diketopiperazine
by an amino group of an oligopeptide is also Definition
possible, which results in an oligopeptide two
amino acids longer. Under standard conditions, dimethyl ether, the
Elongation of an oligopeptide through the simplest ether, is a colorless gas. It is used in
aminolysis of diketopiperazine could play a sig- various chemical syntheses and as an aerosol
nificant role in the prebiotic evolution of propellant. Radio astronomers have found that
oligopeptides, especially in the respect that the dimethyl ether is commonly present in ▶ hot
elongation could take place even in the absence cores, the dense and warm regions of
of guiding templates. In other words, if there ▶ molecular clouds where massive stars are
happens to appear a reaction environment in forming.
Dinitrogen 653
See Also Overview
▶ Hot Core Dinitrogen (N2) refers to ▶ nitrogen occurring in

▶ Molecular Cloud the diatomic form (its most stable form), also
▶ Molecules in Space commonly called molecular nitrogen or nitrogen
gas, in which two nitrogen atoms are linked by a
triple covalent bond (which fills the Lewis elec-
tron rules; the two lone, nonbonding electron
pairs, one on each N atom, lie at a quite low D
Dinitrogen
energy level). N2 is commonly used as a labora-
tory or industrial cryo-coolant, as it has a boiling
point of 196 C and a melting point of 210 C
at atmospheric pressure.
Monoisotopic molecular nitrogen (N2) is
Japan
largely transparent to infrared and visible radia-
tion because it is a homonuclear molecule and has
USA
no electric dipole moment to couple to electro-
magnetic radiation at these wavelengths. Signif-
Washington, DC, USA
icant absorption occurs in the extreme ultraviolet
region, beginning around 100 nm. This absorp-
tion of UV light by N2 is important in the Earth’s
upper atmosphere and the atmospheres of other
planetary bodies.
Keywords
Measurements of the energy required to form
▶ ammonia from N2 and H2 reveal that N2 is
Extreme UV; Photochemistry; Neutral atmo-
unusually stabilized relative to its carbon ana-
sphere; Nitrogen (atomic); Ammonia; Nitrogen
logue, ▶ acetylene, because of the strength of
cycle
the nitrogen-nitrogen triple covalent bond in N2.
This feature dominates the chemistry of
dinitrogen, which is very unreactive at
Synonyms standard temperature and pressure, thus making
it (despite its abundance on Earth) a difficult-to-
Molecular nitrogen; N2; Nitrogen gas fix source of elemental nitrogen for living
organisms.
The conversion of dinitrogen into biologically
Definition useful compounds (usually having N at an oxida-
tion state other than zero) is called “▶ nitrogen
Dinitrogen is a chemical compound formed from fixation,” which is extremely important in both
the covalent bonding of two nitrogen atoms. It is a biological and abiological nitrogen cycling. Bio-
colorless, odorless gas at room temperature and logical nitrogen fixation by bacteria in the root
pressure, which makes up approximately 78 % of nodules of plants produces ammonia (Leigh and
the Earth’s atmosphere (by volume). It is also the Dodsworth 2007). This N2 to NH3 reduction
major component of the atmosphere of ▶ Titan (which is by far the major natural nitrogen source
(one of Saturn’s moons). Dinitrogen exhibits for currently living organisms) is catalyzed by
remarkable stability and is remarkably chemi- nitrogenase enzymes, the active sites of which
cally inert. Liquid dinitrogen is widely used as a contain Fe and Mo atoms, using energy derived
cryo-coolant (b.p. 196 C at atmospheric from the hydrolysis of adenosine triphosphate
pressure). and requiring anoxic conditions.
654 Dione
Abiotic nitrogen fixation occurs in nature

through lightning in the atmosphere, which pro- Dione
duces NOx species (Hill et al. 1980), thanks to very
high temperatures produced in the immediate vicin- Therese Encrenaz
ity of lightning bolts, followed by rapid thermal LESIA, Observatoire de Paris - Section de
quenching that traps high-energy species, or through Meudon, Meudon, France
photocatalysis over various metal-oxide-containing
minerals (Schrauzer et al. 1983). For comparison,
uncatalyzed N2 photochemistry in the Earth’s upper Definition
atmosphere is much less efficient. Precipitation
often contains substantial quantities of ammonium Dione, discovered in 1684 by Giovanni
and nitrate, much of it produced by lightning and Domenico ▶ Cassini, is an icy satellite of
other atmospheric electric phenomena. ▶ Saturn. Its distance to Saturn is 377,000 km
Decay of organisms and their waste products (or 6.2 planetary radii) and its diameter is
results in the formation of nitrate, which eventu- 1,120 km. Its density is 1.5 g/cm3, which indi-
ally may return to the atmosphere as dinitrogen cates a silicate and ice composition. Dione shows
through the microbial reduction of nitrate in a a large variety of structures, with cratered ter-
process known as “denitrification.” rains, faults, and valleys, and albedo contrasts,
Dinitrogen is thought to be a major reservoir indicating an early resurfacing and some internal
of N in interstellar ▶ molecular clouds, although past activity. The longest fissure, Palatine
it cannot be directly detected in these cold regions. Chasma, is almost 400 km long and up to 8 km
Instead, its abundance is deduced from observations wide. A small satellite, Helene, is co-orbiting
of its protonated derivative, N2H+, which has strong with Dione, located at its leading ▶ Lagrangian
pure rotational transitions in the 3-mm region of the point (i.e., Helene is a Trojan moon with respect
spectrum. Besides the Earth, Saturn’s moon Titan is to Dione).
the only other known planetary body with a dense
atmosphere mostly made of dinitrogen, the origin
of which is not yet fully understood.
See Also
See Also ▶ Saturn
▶ Ammonia
▶ Molecular Cloud
▶ Nitrogen
▶ Nitrogen Fixation Dioxygen
▶ Star Formation, Observations
Hiroshi Ohmoto
NASA Astrobiology Institute and Department of
References and Further Reading Geosciences, The Pennsylvania State University,
University Park, PA, USA
Hill RD, Rinker RG, Dale Wilson H (1980) Atmospheric
nitrogen fixation by lightning. J Atmos Sci
37(1):179–192
Leigh JA, Dodsworth JA (2007) Nitrogen regulation in Keywords
bacteria and archaea. Annu Rev Microbiol 61:349–377
Schrauzer GN, Strampach N, Hui LN, Palmer MR, Salehi
Atmospheric oxygen; Evolution of the atmo-
J (1983) Nitrogen photoreduction on desert sands
under sterile conditions. Proc Natl Acad Sci sphere; Oxygen geochemical cycle; Oxygenic
80(12):3873–3876 photosynthesis
Dioxygen 655
Synonyms terrestrial organisms have not contributed to the

long-term (>1,000 year) accumulation of atmo-
Molecular oxygen; O2; Oxygen (molecule) spheric O2. The long-term input of O2 to the
atmosphere has been attributed mostly to the
burial of organic matter (and some to the burial
Definition of pyrite) in marine sediments, which prevents
the reverse Reaction (3) and causes the accumu-
Dioxygen is a molecule formed by the covalent lation of O2 in the atmosphere. About 0.3 % of the
binding of two oxygen atoms. At standard tem- total organic C produced in the oceans is buried in D
perature and pressure, it occurs as a stable gas. O2 sediments to provide about 1013 moles/year of O2
condenses at 90.20 K and freezes at 54.36 K. as the long-term atmospheric input (Holland
1978; Lasaga and Ohmoto 2002). The long-term
output (consumption) of atmospheric O2 has been
Overview carried out primarily by (a) the oxidation of
reducing volcanic gases (e.g., H2, ▶ carbon mon-
The present atmospheric level (PAL) of molecu- oxide (CO), ▶ methane (CH4), ▶ hydrogen sul-
lar oxygen (O2) is 0.209 atm, corresponding to fide (H2S), and sulfur dioxide (SO2)) and (b) the
3.7 1019 moles of O2 in the atmosphere weathering of reduced compounds in rocks (e.g.,
(Holland 1978). The dissociation of water organic C, ferrous ▶ iron, and reduced ▶ sulfur)
(H2O) in the atmosphere and shallow ▶ water (Lasaga and Ohmoto 2002; Kasting 1987).
by UV light is one source of atmospheric O2: Today, the long-term fluxes of O2 production
and consumption appear to be balanced
2H2 O þ hn ! 2H2 þ O2 (1) (Holland 1978; Lasaga and Ohmoto 2002).
The atmospheric pO2 level before the emer-
If hydrogen (H2) can easily escape into space, gence of cyanobacteria at time T1 has been esti-
such as on Mars where the gravity is much less mated to be 1012 PAL, in which the reducing
than on Earth, O2 can continue to accumulate in gases (H2 and CH4) were more abundant than O2
the atmosphere by Reaction (1) alone. (Kasting 1987). O2 became more abundant than
On Earth, a significant production of O2 requires reduced gases at pO2 > 105 PAL (Pavlov and
a continuous consumption of H2 in the reduction of Kasting 2002) at T2, the photic zone became suit-
▶ carbon dioxide (CO2) by organisms: able environments for aerobic organisms and
eukaryotes at pO2 > 5 % PAL (Jahnke and
CO2 þ 2H2 ! CH2 Oðorganic matterÞ þ H2 O Klein 1983) at T3, and deep (as well as shallow)
(2) oceans became oxygenated at pO2 > 50 % PAL
(Kasting 1987; Sarmiento 1992) at T4 (see Table 1).
When Reactions (1) and (2) occur simulta- The theory accepted by most astrobiologists
neously within cells of organisms in the photic postulates the ages of 2.7 Ga (T1), 2.4 Ga (T2
zone, such as ▶ cyanobacteria, ▶ algae, and plants, and T3), and 0.6 Ga (T4), and a subsequent
it is better known as oxygenic ▶ photosynthesis: gradual rise of atmospheric pO2 to the present
level (Kasting 1987; Pavlov and Kasting 2002;
CO2 þ H2 O þ hn ! CH2 O þ O2 (3) Anbar and Knoll 2002). However, some mineral-
ogical, geochemical, and paleontological obser-
Currently, terrestrial and marine organisms vations (Ohmoto 2004; Hoashi et al. 2009)
each produce about 3 1015 moles/year of suggest that the placements of T1–T4 may have
O2 (and also of organic C) (Holland 1978). Essen- occurred before 3.5 Ga and that the atmospheric
tially, all of the organic carbon and O2 produced pO2 level has been maintained within a range of
by terrestrial organisms is converted back to CO2 0.6–2 PAL since T1 by a series of negative feed-
and H2O by the reverse of Reaction (3). Thus, back mechanisms (Lasaga and Ohmoto 2002).
656 Diphosphate
Dioxygen, Table 1 Theories for the oxygenation of the atmosphere and oceans
Time T1 T2 T3 T4
Atmospheric pO2 Level

(PAL) ~10−12 ~10−5 ~5% ~50%
Oxygenic
Photoautotrophs
Oxic Atmosphere
(pO2 > pH2)
Oxic Photic Zone
Oxic Deep Oceans
Time before Present (Ga)

~2.7 ~2.4 ~2.4 ~0.6
(Theory 1)
Time before Present (Ga)

>3.5
(Theory 2)

Diphosphate
Anbar AD, Knoll AH (2002) Proterozoic ocean chemistry
and evolution: a bioinorganic bridge? Science ▶ Pyrophosphate
297:1137–1142
Hoashi H et al (2009) Primary heaematite formation in an
oxygenated sea 3.46 billion years ago. Nat Geosci
2:301–306
Holland HD (1978) The chemistry of the atmosphere and
Diplogen
oceans. Wiley, New York
Jahnke L, Klein HP (1983) Effects of low levels of ▶ Deuterium
oxygen on Sacchromyces cerevisial. Orig Life
9:329–334
Kasting J (1987) Theoretical constraints on
oxygen and carbon dioxide concentrations in the
Precambrian atmosphere. Precambrian Res Dirac, Paul
34:205–229
Lasaga AC, Ohmoto H (2002) The oxygen geochemical Fernando B. Figueiredo
cycle: dynamics and stability. Geochim Cosmochim
CITEUC, University of Coimbra, Coimbra,
Acta 66:361–381
Ohmoto H (2004) Archean atmosphere, hydrosphere and Portugal
biosphere. In: Ericksson PG et al (eds) The Precam-
brian Earth: tempos and events. Elsevier, Amsterdum,
pp 361–388
Pavlov AA, Kasting J (2002) Mass-independent fraction-
History
ation of sulfur isotopes in Archean sediments: strong
evidence for an anoxic Archean atmosphere. Astrobi- Paul Adrien Maurice Dirac (1902–1984) was a
ology 2:27–41 British quantum physicist awarded in 1933 with
Sarmiento J (1992) Biogeochemical ocean
the Nobel Prize (which he shared with
models. In: Trenberth KE (ed) Climate system
modeling. Cambridge University Press, Cambridge, Schrödinger) for the “discovery of new produc-
pp 5–59 tive forms of atomic theory.”
Direct-Imaging, Planets 657
Dirac established, independently of Fermi, the and the atmosphere. The challenge of developing
statistical laws (now called Fermi-Dirac statis- these instruments – ▶ coronagraphs and interfer-
tics) that ruled fermions, particles systems sub- ometers with ▶ adaptive optics systems – has
ject to the Pauli exclusion principle. His scientific caused direct imaging to lag behind other
work led to introduce the concept of antimatter, exoplanet detection techniques. But direct imag-
which plays a central role in modern particle ing offers access to planets with host stars and
physics and cosmology. orbital properties that make them unreachable by
In 1932 Dirac was appointed Lucasian profes- other techniques, and a means to find Earth-like
sor of mathematics at the University of planets around nearby stars and characterize their D
Cambridge, which was also Newton’s chair. atmospheres. The few planets that have been
Stephen Hawking was Dirac’s successor in the detected by direct imaging so far tend to be hot,
Lucasian chair. young giant planets, located far (>20 AU) from
their host stars. The presence of such objects
presents a challenge for some theories of planet
See Also formation.
▶ Fermi, Enrico
History
In 2005, Chauvin et al. (2005) imaged a ~5

Directed Evolution Jupiter-mass companion to a nearby ▶ brown
dwarf using the ▶ VLT/NACO. Imaged planets
▶ Evolution, In Vitro have also been reported in the HR 8799 and
▶ Evolution, Molecular Fomalhaut systems, though the masses of these
objects remain uncertain and some of them may
turn out to be in the brown dwarf, rather than
planet, mass range. The directly imaged planet
with the closest orbit (8 AU) to its star has been
Direct-Imaging, Planets
reported in the Beta Pic system by Lagrange
et al. (2010).
Marc Kuchner
NASA Goddard Space Flight Center, Exoplanets
and Stellar Astrophysics Laboratory, Greenbelt,
See Also
MD, USA
▶ Adaptive Optics
▶ Brown Dwarf
Definition
▶ Coronagraphy
Direct imaging of planets refers to the effort to
▶ Interferometry
detect and study ▶ exoplanets from the light
▶ VLT
emitted or scattered by the planets themselves,
as opposed to inferring the existence and proper-
ties of planets from their effects on the images or References and Further Reading
spectra of the stars they orbit. Directly ▶ imaging
an exoplanet, i.e., collecting light from the planet Chauvin G et al (2005) Giant planet companion to
2MASSW J1207334-393254. Astron Astrophys 438:
and distinguishing it from light from the host star,
L25–L28
requires special instruments to overcome distor- Lagrange A-M et al (2010) A giant planet imaged in the
tions caused by conventional ▶ telescope optics disk of the young star b pictoris. Science 329:57–59
658 Disinfection
gravitational instabilities in the ▶ protoplanetary

Disinfection disk. Numerical simulations have shown that
clumps of about one to several Jupiter masses
Catharine A. Conley can form in sufficiently gravitationally unstable
NASA Headquarters, Washington, DC, USA disks. Protoplanetary disks can become gravita-
tionally unstable if they are sufficiently cool
and/or massive. The condition for gravitational
Definition
instability can be determined from the value of
the Toomre parameter Q. Protoplanetary disks
Disinfection is an action taken to obtain the reduc-
will be unstable for Q < ~1. Once a local insta-
tion of the number of viable microorganisms pre-
bility occurs, a gravitationally bound
sent in an environment (gas, liquid, surface, etc.).
sub-condensation region (clump) is created. If
the cooling time in this region of the disk is
See Also shorter than (or comparable to) the dynamical
time, the fragment contracts, and eventually
▶ Bioburden Reduction
evolves to become a giant planet.
▶ Inactivation
▶ Pasteurization
▶ Sterilization See Also
▶ Gravitational Instability
Disk Instability, Model for Giant ▶ Q (Toomre Parameter)
Planet Formation
Yann Alibert1 and Ravit Helled2

1
Space Research and Planetary Sciences, Physics Dismutation
Institute, University of Bern, Bern, Swiss
2
Geophysical, Atmospheric and Planetary ▶ Disproportionation
Sciences, Tel Aviv University, Raymond and
Beverly Sackler Faculty of Exact Scienes,
Tel Aviv, Israel Disproportionation

Keywords Earth–Life Science Institute (ELSI), Tokyo
Planet formation; Protoplanetary disk; Japan
Q (Toomre parameter) Institute for Advanced Study, Princeton, NJ,
USA
Synonyms Blue Marble Space Institute of Science,
Washington, DC, USA
Gravitational instability (Model for Giant Planet Center for Chemical Evolution, Georgia Institute
Formation) of Technology, Atlanta, GA, USA
Definition
Synonyms
The disk instability model suggests that ▶ gas
giant planets form directly as a result of (local) Dismutation
Dissolved Inorganic Carbon Equilibrium 659
Definition electron acceptor must be reduced to guaranty the

generation of sufficient energy, and the product is
In chemistry, a disproportionation reaction is a excreted into the environment, while in the
type of chemical reaction in which two identical ▶ assimilative metabolism, only enough amount
compounds react to form two or more new com- of the compound is reduced to satisfy the needs
pounds of a dissimilar type. Usually, these involve for cell growth, and the products are normally
reaction compounds containing an element in the converted into cell material.
same oxidation state reacting to form two com-
pounds containing that element but with both D
higher and lower oxidation states, for example, in
the Cannizzaro reaction with formaldehyde: See Also
2 HCHO ! HCOOH þ CH3 OH (1) ▶ Anaerobic Respiration

▶ Assimilative Metabolism
An example of a disproportionation reaction in ▶ Electron Acceptor
which no redox chemistry takes place is the ion- ▶ Metabolism
ization of water: ▶ Reduction
▶ Sulfate Reducers
2H2 O ! H3 Oþ þ OH (2)
See Also
Dissolved Inorganic Carbon
▶ Fractionation
Equilibrium
▶ Redox Potential
Dissimilative Metabolism Japan
Juli Peretó USA
Institut Cavanilles de Biodiversitat i Biologia Blue Marble Space Institute of Science,
Evolutiva, Universitat de València, València, Washington, DC, USA
Spain Center for Chemical Evolution, Georgia Institute
Definition
Definition
Dissimilative ▶ metabolism is the process by
which an inorganic compound (NO3, SO42, The dissolved inorganic carbon equilibrium is the
Fe3+, CO2) is reduced because it is used as an equilibrium between CO2 (g), HCO3 (aq),
▶ electron acceptor for ▶ anaerobic respiration. H2CO3 (aq), and CO32 (aq) when CO2 is
Dissimilative metabolism is conceptually very dissolved in water. The equilibrium is dependent
different from the ▶ assimilative metabolism by on the pressure of the gas, the overlying head-
which the same compounds are reduced for its space and gas partial pressure, the solution tem-
use as nutrient source. There are important dif- perature, the ▶ pH of the solution, and the
ferences between both types of metabolism. In concentration of various solutes such as electro-
the dissimilative metabolism, a large amount of lytes. It is significant because dissolved carbonate
660 Distal Impact Ejecta
species may have a significant impact on the pH See Also

of natural waters:
CO2 ðgÞ $ CO2 ðaqÞ (1) ▶ Isotopic Fractionation (Planetary Process)
CO2 ðaqÞ þ H2 O $ H2 CO3 ðaqÞ

$ HCO3 ðaqÞ þ Hþ $ CO2 ðaqÞ þ 2Hþ
(2) Disulfide Bond
See Also Kensei Kobayashi
▶ Carbonate Hodogaya-ku, Yokohama, Japan
▶ pH
Synonyms
Distal Impact Ejecta SS-bond
▶ Spherules
Definition
A disulfide bond is a covalent bond between two

sulfur atoms (-S-S-) formed by the coupling of
Distillation, Rayleigh
two ▶ thiol (-SH) groups. ▶ Cysteine, one of the
20 protein amino acids, has a thiol group in its
Francis Albarède
side chain and can easily be dimerized to ▶ cys-
tine in aqueous solution by forming a disulfide
France
bond. In many protein molecules, disulfide bonds
between cysteine residues are essential for pro-
tein folding. Disulfide bonds in proteins are
Definition
cleaved by heating or by the addition of reducing
reagents, which leads to protein denaturation.
This term applies to chemical and isotopic
▶ fractionation during phase transformation
when one phase progressively evolves from
another of limited size and is continuously See Also
removed. Upon evaporation of an alcoholic solu-
tion, ethanol concentrates into the first vapor ▶ Amino Acid
extracts and, as a result of the finite original sup- ▶ Cysteine
ply, becomes depleted in the residual liquid. Dis- ▶ Cystine
tillation, which bears the name of “Rayleigh ▶ Protein
distillation” from Lord Rayleigh who provided ▶ Thiol
the original equation, greatly enhances fraction-
ation over exchange at equilibrium. Distillation as
described for spirit amounts to fractional evapora-
tion: changing phases leads to fractional conden- Diversity of Life
sation (rain), fractional crystallization (magmatic
differentiation), fractional melting, etc. ▶ Biodiversity
DLR, Germany 661
Definition
Division
In 1984, the Deutsche Agentur f€ur Raumfahrtan-
▶ Phylum gelegenheiten (DARA) – German Aerospace
Agency – was created. It was commissioned to
oversee the planning and coordination of state-
Dixon Island Formation, Western funded German aerospace initiatives and to rep-
Australia resent Germany at meetings of international bod-
ies. In the same year the DFVLR is renamed the D
Nicholas Arndt Deutsche Forschungsanstalt f€ur Luft-und
ISTerre, Université Grenoble Alpes, France Raumfahrt (DLR) – the German Aerospace Cen-
ter. The present DLR was established in 1997 by
Definition the amalgamation of these two bodies, which
henceforth will be called the Deutsches Zentrum
The Dixon Island Formation is a 3.2-Ga-old f€ur Luft- und Raumfahrt – German Aerospace
sequence of lightly metamorphosed but highly silic- Center. Within all these years, Germany was
ified cherts, organic-rich black shales and tuffs able to implement more than 100 space missions
belonging to the Cleaverville Group and located in both nationally and within the framework of
the ▶ Pilbara Craton, Western Australia. The unit is international cooperation.
thought to have formed through shallow marine The space activities include Germany’s
hydrothermal alteration of Archean ▶ oceanic national space program, DLR’s “space” research
crust. Possible microbial mats and bacteria-like and development program, and Germany’s con-
structures have been observed and are considered tributions to the ▶ European Space Agency
by some to be traces of early life. The sequence was (ESA) as well as the European Organization for
the target of the Dixon Island-Cleaverville Drilling the Exploitation of Meteorological Satellites
Project (DXCL-DP). The most important objective (EUMETSAT). Germany is the first contributor
of the DXCL-DP was to understand the nature of the to the ESA mandatory program and the second in
Middle Archean marine environment, influenced by the optional programs. In addition, the agency
hydrothermal activity, through detailed and system- designs and supervises space and security
atic study of fresh drill core samples. research projects under the sixth and seventh
EU research framework program.
See Also DLR’s mission comprises research aimed at
protecting the environment; development of envi-
▶ Archean Drilling Projects ronmentally friendly technologies to promote
▶ Biomarkers mobility, communication, and security; as well
▶ Pilbara Craton as exploration of the Earth and the Solar System
and space, which leads to new knowledge about
the origin and development of the Solar System,
its planets, and, hence, about the emergence of life.
DLR, Germany The center has 29 institutes and facilities
at 13 locations in Germany: Berlin, Bonn, Braun-
Michel Viso schweig, Bremen, Cologne (headquarters),
CNES/DSP/SME, Vétérinaire/DVM, Goettingen, Hamburg, Lampoldshausen,
Astro/Exobiology, Paris Cedex 1, France Neustrelitz, Oberpfaffenhofen, Stuttgart, Trauen,
and Weilheim. Approximately 6,500 persons are
Synonyms currently working for DLR.
For further information, see http://www.
German Aerospace Center dlr.de.
662 DNA
Avery, Colin MacLeod, and Maclyn McCarty

DNA demonstrated that DNA taken from a disease-
causing strain of the bacterium Streptococcus
Irena Mamajanov and Nicholas V. Hud pneumoniae permanently transformed a
School of Chemistry and Biochemistry, Georgia non-virulent form of the organism into a virulent
Institute of Technology, Atlanta, GA, USA one (Avery et al. 1944). Additional evidence was
provided in 1951 by Alfred Hershey and Martha
Chase, who demonstrated that when the bacteri-
Keywords ophage T2 infects Escherichia coli, it is the DNA
of the bacteriophage, not its protein coat, which
Adenine; Adenosine; Cytidine; Cytosine; Deoxy- enters the host cell and provides the genetic infor-
ribose; DNA; Guanine; Guanosine; Phosphate; mation for bacteriophage replication (Hershey
Purine; Pyrimidine; Ribose; RNA; Thymidine; and Chase 1951).
Thymine The double-helix structure of DNA was first
proposed by James D. Watson and Francis Crick
in 1953 (Watson and Crick 1953). The Watson-
Synonyms Crick model was largely based on X-ray fiber
diffraction data of DNA obtained by Rosalind
Deoxyribonucleic acid; Polydeoxyribonucleic Franklin and Raymond Gosling (Franklin and
acid Gosling 1953) and subsequent diffraction pat-
terns obtained by Maurice Wilkins and
coworkers (Wilkins et al. 1953). Biochemical
Definition information obtained by Erwin Chargaff regard-
ing the 1:1 stoichiometry of G:C and A:T in DNA
DNA (deoxyribonucleic acid) is the polymer extracted from a wide range of organisms also
that stores the genetic information of all facilitated Watson and Crick’s elucidation of the
living cells and many viruses. DNA is a linear double-helix structure (Elson and Chargaff
polymer of phosphodiester-linked 20 -deoxyribose 1954). Crick, Watson, and Wilkins shared the
sugars with heterocyclic bases (nucleobases) 1962 Nobel Prize in Physiology and Medicine
attached to the 10 carbon of the 20 -deoxyribose for their contributions to determining the struc-
sugars. The four nucleobases of DNA are ture of the DNA double helix.
adenine (A), guanine (G), cytosine (C), and
▶ thymine (T). DNA within a living cell DNA Structure
primarily exists in the form of a double helix in The primary structure of DNA is generally
which two polymer strands are held together by described in terms of nucleotides, the molecular
Watson-Crick base pairs (i.e., AT and GC unit defined as a nucleobase connected by a gly-
base pairs). cosidic bond at the 10 position of a phosphory-
lated 20 -deoxyribose sugar. Along the backbone
of a DNA polymer, or strand, nucleosides are
Overview and Key Research Findings connected to each other through the phosphate
groups. A phosphodiester linkage exists between
DNA was discovered by Johann Friedrich the 30 position of each ▶ nucleoside and the 50
Miescher in 1863. He separated the so-called position of the next nucleoside along a given
nuclein from cells and showed that it was a pre- DNA strand (Fig. 1). The order of deoxyribose
viously uncharacterized phosphorus-containing nucleotides (dA, dG, dC, and dT) along a strand is
material and hypothesized its involvement in the chemical basis for coding the genetic infor-
heredity (Dahm 2005). It was not until 1944 that mation contained within a DNA polymer.
this hypothesis was confirmed, when Oswald A single ▶ nucleotide that is not phosphorylated,
DNA 663
DNA, Fig. 1 The chemical ge

structure of DNA which ed
ve
illustrates Watson-Crick g roo
jor O
base pairing and the O ma H H
O
directionality of the N N
T
N
deoxyribose- −O P O H 3'
N
phosphodiester backbone N O
O A O
e
N edg O
O N
o o ve O−
5' 1' g r H P
5' 4' o r g e
min ed O
3' 2'
roo
ve H N O D
g C
O a jor O 5'
m N N
− O N
O P H
O
GN O O
N H O
3' O N N
g e O
d P
H
o v ee O
ro
or g O
O min
that is, only the base connected to the sugar, is

referred to as a nucleoside.
The repeating 30 -phosphate-50 linkages
between nucleotides provide the polymer direc-
tionality for DNA that is inseparable from its
synthesis and function. For example, DNA is
synthesized by natural polymerases in the 50 -30
direction. That is, a DNA polymer being
extended by a polymerase will have new nucleo-
tides added to the terminal nucleotide at the free
30 -OH group.
The secondary structure of DNA within living
organisms is predominantly a double helix
formed by two strands with complementary
nucleotide sequences, that is, sequences that sup-
port Watson-Crick base pairing along their entire DNA, Fig. 2 A space-filling model of a 12 base-pair
duplex of B-form DNA. (Left) The model orientated
lengths. The two strands of a Watson-Crick dou- such that reader is looking into the major groove at the
ble helix run in opposite directions (as defined by midpoint of the duplex. (Right) The same model, but
the directionality of their polymer backbones) orientated such that the reader is looking into the minor
(Fig. 1). A Watson-Crick duplex is therefore groove at the midpoint of the duplex. Backbone atoms are
shown in white, base carbon atoms in green, base nitrogen
said to have an antiparallel arrangement of its atoms in blue, and base oxygen atoms in red. For clarity,
two strands. While complementary hydrogen hydrogen atoms are not shown
bonding between nucleobases (i.e., A with T;
G with C) provides specificity for base-pair for-
mation, interstrand nucleobase stacking interac- The predominant helical structure of DNA
tions apparently provide the most significant within a cell and in vitro at high humidity (e.g.,
energetic contribution to helix stability above 95 % relative humidity) is known as the
(Yakovchuk et al. 2006). B-form helix (Fig. 2). The B-form helix is
664 DNA
approximately 2 nm wide with a right-handed sequences without direct contact between the
twist and a helical rise of around 3.4 nm per ten protein and the DNA bases that differ between
nucleotides. One complete turn of the B-form two otherwise identical DNA sequences. Such
helix therefore consists of 10–10.5 base pairs. protein-DNA recognition is referred to as indirect
The B-form helix has two grooves that are of readout and can have its physical origins in the
different widths and depths (Fig. 2). The wider flexibility of a nucleic acid sequence, water-
groove is referred to as the major groove and the mediated contacts, and sequence-specific cation
narrower one as the minor groove. localization (Watkins et al. 2010).
Under low humidity/dehydrating conditions,
duplex DNA will undergo a structural transition DNA Packaging and Supercoiling
from the B-form to the A-form helix (Saenger The amount of DNA necessary for coding all
1984), which is the same helical structure associ- aspects of a living organism has necessitated the
ated with duplex ▶ RNA. The A-form helix is evolution of highly efficient DNA packaging
also right handed but with a helical repeat of mechanisms. For example, the human ▶ genome
approximately 11 base pairs per helical turn. contains almost three billion base pairs of DNA
The width and depth of the major and minor per haploid genome. If the DNA contained within
grooves of the A-form helix also differ signifi- the diploid genome of a human cell was fully
cantly from the B-form helix. extended, it would have a linear measurement of
nearly 2 m. Given that the diameter of a typical
DNA Coding of Proteins human cell nucleus is around 2 mm, genomic
The genetic information that codes for the amino DNA must be packaged in a highly condensed
acid sequence of a particular protein is tran- state. In eukaryotic cells, the packaging of DNA
scribed from a DNA polymer by a is accomplished by a group of evolutionarily
DNA-directed RNA polymerase into messenger conserved and positively charged proteins that
RNA (or mRNA). The information contained in form the histones, octomeric protein assemblies
the mRNA is then translated into a protein by the that are each wound by 145 base pairs of DNA
▶ ribosome, tRNA molecules, and other proteins (Luger et al. 1997).
necessary for protein synthesis and the overall The genomic DNA of bacterial cells is pack-
process of translation (Kornberg 2007). aged by a combination of molecular and physical
The transcription of many genes in eukaryotic factors, including supercoiling, molecular
cells and bacteria is regulated by the binding of crowding, polyamines, and protein binding
proteins known as activators and transcription (Zimmerman 2006). Supercoiling refers to the
factors to specific DNA sequences that are over twisting of a topologically constrained
located “upstream” from the protein-coding DNA duplex (a covalently closed circle of duplex
region of a gene (Kornberg 2007). These gene- DNA is the simplest example of a topologically
regulating proteins often recognize their target constrained DNA molecule). Much like a rubber
sequence through complementary hydrogen- band that has been rolled and twisted, a
bonding interactions between amino acid side supercoiled and topologically constrained DNA
chains and the edges of the nucleobases that are molecule will tend to collapse onto itself. Molec-
accessible in the major and minor grooves of ular crowding agents and positively charged
duplex DNA. The major groove appears to be polyamines also promote the collapse of DNA
the preferred groove for sequence recognition into a highly condensed state within bacterial
by DNA-binding proteins due to the greater dis- cells by promoting the general phenomenon of
similarity between the pattern of hydrogen donor DNA condensation (i.e., the propensity for DNA
and acceptor groups of the major groove base to interact with itself in preference to the sur-
edges, in comparison to the minor groove rounding solvent) (Bloomfield 1996). It appears
(Seeman et al. 1976). Some transcription factors that two highly covered bacterial proteins known
are able to differentiate between similar DNA as HU and IHF facilitate DNA packaging by
DNA 665
promoting local bends in the DNA double helix exist between phosphate groups across the
(Sarkar et al. 2007). major and minor grooves. The screening of
these repulsions by cations is reflected by the
DNA Triplexes and Quadruplexes observed dependence of DNA duplex stability
DNA double helix formed between a homopurine on the concentration of salt in a solution, as well
strand (i.e., only A and G nucleobases) and a as the valency of the cation (e.g., monovalent
homopyrimidine strand (i.e., only T and versus divalent). As mentioned above, the role
C nucleobases) under certain conditions (e.g., that hydration plays in regulating DNA helical
high salt or solutions containing Mg2+) can form structure is illustrated by the B- to A-form struc- D
a triplex secondary structure with a third strand tural transition that occurs below 95 % relative
bound in the major groove (Frank-Kamenetskii humidity.
and Mirkin 1995). This third strand forms the Although it had long been postulated that
so-called Hoogsteen pairs with the major groove DNA secondary structures have an absolute
edges of the ▶ purine bases of the Watson-Crick requirement for water, recent studies show that
duplex. There are two possible triplexes: pyrim- water is not the only media able to sustain native
idine triplexes, in which the third strand is com- DNA structures. A new class of water-free deep
posed only of pyrimidine nucleobases, and purine eutectic solvents has been reported to sustain
triplexes, in which the third strand is composed duplex, triplex, and G-quadruplex structures
only of purine nucleobases. In the case of pyrim- (Mamajanov et al. 2010).
idine triplexes, the C nucleobases require proton-
ation in order to form a Hoogsteen pair with the The Origin of DNA
G bases. Because the pKa of the C nucleobase is The discovery of catalytic RNA molecules, or
around 4.5, pyrimidine triplexes are most stable ribozymes, and the ever-increasing evidence
under slightly acidic conditions. Triplex-forming that RNA is the central polymer of life have
nucleotides have been the object of intensive created great enthusiasm for the hypothesis that
study as potential therapeutic agents for the mod- RNA once carried out both the information stor-
ulation of gene expression activity in vivo age needs are now satisfied by DNA as well as the
(Frank-Kamenetskii and Mirkin 1995). catalytic tasks now accomplished by protein
G-rich DNA sequences (e.g., TGGGGT) are enzymes. There are additional reasons to suspect
capable of forming four-stranded secondary that RNA preceded DNA (Dworkin et al. 2003),
structures called G-quadruplexes that are held including the appearance of ▶ ribose in model
together by G-tetrads in which the prebiotic reactions and the presumed lack of a
G nucleobases are in a planar, cyclic Hoogsteen prebiotic route to deoxyribose; activated ribonu-
hydrogen-bonded arrangement (Davis 2004). cleotides polymerize more readily than activated
These structures are further stabilized by cation deoxyribonucleotides; the biosynthesis of deoxy-
coordination by the guanine O6 carbonyl groups ribonucleotides is through ribonucleotides in
in the center of the G-tetrad. G-quadruplexes are extant life. Ribosides are subunits of many coen-
potentially formed in vivo by a large number of zymes, some of which have prebiotic syntheses.
G-rich DNA sequences within the human The much greater chemical stability of DNA
genome (Huppert and Balasubramanian 2007), under conditions of neutral and basic pH, as com-
which has been taken as evidence that pared to RNA, would have provided a strong
G-quadruplexes are involved gene regulation. evolutionary advantage for any organism that
developed a means to synthesize DNA for use
DNA Interactions with Water and Cations in long-term information storage. In contempo-
DNA-water and DNA-cation interactions are rary life, ▶ ribonucleotide reductases are respon-
intimately associated with DNA structure. The sible for converting ribonucleotides into
high dielectric constant of water and cations deoxyribonucleotides through a free radical-
screen the electrostatic repulsions that would mediated process. The lack of known ribozymes
666 DNA
that can carry out such reactions has been an anhydrous solvent (Mamajanov et al. 2010)
suggested as evidence that DNA must have supports the proposal that nucleic acids might
appeared only after coded proteins were part of have first evolved in nonaqueous or mixed
life (Dworkin et al. 2003). However, it has more aqueous-organic media.
recently been argued that RNA could have syn-
thesized DNA by a nonreductive mechanism
(e.g., via aldol condensation chemistry), which See Also
could have allowed DNA to appear before coded
proteins (Burton and Lehman 2009). ▶ Adenine
▶ Cytosine
▶ DNA Damage
Basic Methodology ▶ DNA Repair
▶ DNA Sequencing
See ▶ Nucleic Acids ▶ Double Helix
▶ Electrophoresis
▶ Extraterrestrial Delivery of Organic
Applications Compounds
▶ Genetics
Genetic engineering, the artificial manipulation ▶ Genome
of genetic material for the production of natural ▶ Guanine
and novel proteins, is a major branch of biotech- ▶ Nucleic Acid Base
nology that would not exist without our funda- ▶ Nucleic Acids
mental understanding of DNA structure and its ▶ Nucleoside
use within living cells. Examples of genetic engi- ▶ Nucleotide
neering applications in medicine include bacteria ▶ Oligonucleotide
that produce human insulin and proteins for viral ▶ Phylogenetic Tree
vaccines. In addition, DNA research has given ▶ Phylogeny
rise to bioinformatics, a field that utilizes infor- ▶ Polymerase Chain Reaction
mation technology, from data storage to DNA ▶ Prebiotic Chemistry
sequence analysis, to provide insights regarding ▶ Purine Bases
biological evolution, from recent events to the ▶ Pyrimidine Base
root of the tree of life (Garrett and Grisham ▶ Replication (Genetics)
2004). Finally, DNA is now routinely used in ▶ Ribonucleoside
forensic science, with the help of PCR and bioin- ▶ Ribonucleotide
formatics, to identify suspects, as well as victims, ▶ Ribose
and to provide what can be the most critical ▶ Ribosome
evidence in support of a conviction or acquittal. ▶ RNA
▶ RNA World
▶ Sequence
Future Directions ▶ Thymine (T)
▶ Watson-Crick Pairing
Nucleic acid research is constantly moving for-
ward with deeper insights into the forces that
govern DNA structure and how DNA functions References and Further Reading
within living cells, as well as refinement of
Avery O, MacLeod C, McCarty M (1944) Studies on
biotechnological applications. From the the chemical nature of the substance inducing trans-
astrobiological perspective, the discovery that formation of pneumococcal types. J Exp Med
DNA secondary structure can be maintained in 79:137–157
DNA Damage 667
Bloomfield V (1996) DNA condensation. Curr Opin Struct

Biol 6:334–341 DNA- (or RNA-)Dependent DNA
Burton AS, Lehman N (2009) DNA before proteins?
Recent discoveries in nucleic acid catalysis strengthen Polymerase
the case. Astrobiology 9:125–130
Dahm R (2005) Friedrich Miescher and the discovery of ▶ DNA Polymerase
DNA. Dev Biol 278:274–288
Davis JT (2004) G-quartets 40 years later: from 50 -GMP to
molecular biology and supramolecular chemistry.
Angew Chem Int Ed Engl 43:668–698
Dworkin JP, Lazcano A, Miller SL (2003) The roads to DNA (or RNA-)-Dependent RNA D
and from the RNA world. J Theor Biol 222:127–134 Polymerase
Elson D, Chargaff E (1954) Regularities in the composi-
tion of pentose nucleic acids. Nature 173:1037–1038
Frank-Kamenetskii MD, Mirkin SM (1995) Triplex DNA ▶ RNA Polymerase
structures. Annu Rev Biochem 64:65–95
Franklin RE, Gosling RG (1953) Molecular configuration
in sodium thymonucleate. Nature 171:740–741
Garrett RH, Grisham CM (2004) Biochemistry, 3rd edn.
Brooks Cole, Charlottesville DNA Damage
Hershey AD, Chase M (1951) Genetic recombination and
heterozygosis in bacteriophage. Cold Spring Harb
Symp Quant Biol 16:471–479 Thierry Douki and Jean Cadet
Huppert JL, Balasubramanian S (2007) G-quadruplexes in Laboratoire Lésions des Acides Nucléiques,
promoters throughout the human genome. Nucleic Institute Nanosciences et Cryogénie/CEA,
Acids Res 35:406–413
Grenoble, France
Kornberg RD (2007) The molecular basis of eukaryotic tran-
scription. Proc Natl Acad Sci U S A 104:12955–12961
Luger K, Mader AW, Richmond RK, Sargent DF, Rich-
mond TJ (1997) Crystal structure of the nucleosome core Keywords
particle at 2.8 angstrom resolution. Nature 389:251–260
Mamajanov I, Engelhart A, Bean H, Hud N (2010) DNA
and RNA in anhydrous media: duplex, triplex, and Bipyrimidine photoproducts; Cyclobutane
G-quadruplex secondary structures in a deep eutectic pyrimidine dimers; DNA oxidation products;
solvent. Angew Chem Int Ed 49:6310–6314 DNA strand breaks; 8-oxo-7,8-dihydro-
Saenger W (1984) Principles of nucleic acid structure.
20 -deoxyguanosine; Pyrimidine (6-4) pyrimidone
Springer, New York
Sarkar T, Vitoc I, Mukerji I, Hud NV (2007) Bacterial photoproducts; Spore photoproduct
protein HU dictates the morphology of DNA conden-
sates produced by crowding agents and polyamines.
Nucleic Acids Res 35:951–961
Seeman N, Rosenberg J, Rich A (1976) Sequence-specific
Synonyms
recognition of double helical nucleic acids by proteins.
Proc Natl Acad Sci U S A 73:804–808 Alteration
Watkins D, Mohan S, Koudelka GB, Williams LD
(2010) Sequence recognition of DNA by protein-induced
conformational transitions. J Mol Biol 396:1145–1164
Watson JD, Crick FHC (1953) Molecular structure of Definition
nucleic acids: a structure for deoxyribose nucleic
acid. Nature 171:737–738 ▶ DNA damage consists in chemical modifica-
Wilkins MHF, Stokes AR, Wilson HR (1953) Molecular
tions of the deoxyribonucleic acid components
structure of deoxypentose nucleic acids. Nature
171:738–740 that include alterations of the four main purine
Yakovchuk P, Protozanova E, Frank-Kamenetskii MD (adenine, guanine) and pyrimidine (cytosine,
(2006) Base-stacking and base-pairing contributions thymine) bases, the relatively minor
into thermal stability of the DNA double helix. Nucleic
5-methylcytosine base and the 2-deoxyribose
Acids Res 34:564–574
Zimmerman SB (2006) Shape and compaction of moiety. According to the damaging agents that
Escherichia coli nucleoids. J Struct Biol 156:255–261 may be endogenous (reactive oxygen and
668 DNA Damage
nitrogen species such as hydroxyl radical, inside manned space vessels due to an efficient
peroxynitrite, etc.) and exogenous (solar light, shielding against the UV radiation components
ionizing radiation, alkylating compounds, etc.), involving the vacuum-UV (140< l <200 nm),
several classes of DNA lesions may be generated. UVC (200< l <280 nm), UVB (280< l
These include single- and double-strand breaks, <320 nm), and UVA (320< l <400 nm) pho-
normal and oxidized abasic sites, single modified tons. This protection is much more limited
bases (oxidized lesions, alkylated adducts, addi- against high-charge (Z) and high-energy
tion products with reactive aldehyde arising (E) (HZE) particles of galactic cosmic radiation
from the breakdown of lipid peroxides, and (GCR) that, in addition to the HZE particles (1 %
2-deoxyribose oxidation), tandem modifications of the nucleonic component), also include high-
(intrastrand bipyrimidine photoproducts, vicinal energy protons (87 %) and a-particles (He ions)
oxidized bases), DNA-protein cross-links, (12 %) as well as electrons. Of concern are also
interstrand cross-links, and clustered lesions solar particle radiations that are emitted during
(association of several oxidized bases, single- or solar wind and erratic solar flares, consisting
double-strand breaks produced within one to two mostly of protons with very small amounts of
helix turns by several radiation-induced a-particles and HZE ions (Durante and Cucinotta
radical hits). 2008).
History Basic Methodology
The discovery and characterization of cis-syn Major progress has been made during the last two
cyclobutane thymine dimer at the beginning of decades in the development of accurate and sen-
the 1960s, the main UVB- and UVA-mediated sitive methods aimed at measuring photo- and
degradation product of isolated and cellular radiation-induced damage to cellular DNA. This
DNA, has provided a strong impetus to investi- has benefited from the availability of new meth-
gations in the fields of genotoxicity, mutagenesis, odological approaches including high-
and ▶ DNA repair. performance liquid chromatography coupled
with electrospray ionization-tandem mass spec-
trometry detection (HPLC-ESI-MS/MS). Thus,
Overview the UV-induced DNA photoproducts that mostly
consist of cyclobutane pyrimidine dimers
This entry focuses on the two main classes of (CPDs), pyrimidine (6-4) pyrimidone photoprod-
DNA modifications that may be produced upon ucts (6-4PPs) at the four main bipyrimidine
exposure of living systems to deleterious space sequences together with 5,6-dihydro-5-
conditions. These conditions comprise high vac- (a-thyminyl)-thymine, the so-called spore photo-
uum, wide range of temperature variations, and product (SP), can be measured by HPLC-MS/MS
ionizing and UV radiations that have both galac- (Cadet et al. 2005; Cadet and Douki 2010). Poly-
tic and solar origins (Nicholson et al. 2000). clonal and monoclonal antibodies are also avail-
These effects include DNA photoproducts that able for monitoring the formation of bipyrimidine
are the main alterations generated upon exposure photoproducts in cellular DNA and in tissue in a
to solar extraterrestrial UV radiation of resistant more semiquantitative way. HPLC-MS/MS is the
microorganisms which may be present in outer method of choice for assessing the formation of
space. Under the latter conditions, the contribu- single and clustered oxidatively generated base
tion of the second class of DNA damage that damage in cellular DNA at least under acute
consists of radiation-induced degradation prod- conditions of ionizing radiation exposure (Cadet
ucts involving mostly oxidative reactions is rela- et al. 2010). The modified version of the comet
tively minor. However, the situation is different assay and the alkaline elution technique that both
DNA Damage 669
require the use of DNA repair glycosylases Bacterial Spores

to reveal classes of oxidatively generated The spectrum of DNA photoproducts in UVC- or
damage such as oxidized bases and modified UVB-irradiated bacterial spores is very different
purine bases are suitable alternatives although from what is observed in vegetative cells. Thus,
less specific than HPLC-MS/MS to deal with the spore photoproduct (SP) that is another
the detection of low amounts of radiation- bithymine lesion is formed almost exclusively at
induced DNA damage. the expense of CPDs and 6-4PPs in Bacillus
subtilis spores (Cadet and Douki 2010). The pro-
nounced changes in DNA photoreactivity have D
Key Research Findings been accounted for by notable structural modifi-
cations of DNA molecules that were shown to
DNA Photoproducts adopt an A-like conformation. This was found to
The UVC and UVB ▶ photochemistry of DNA result from the binding of small acid-soluble pro-
that is triggered by the direct excitation of the teins (SASP) in a dehydrated environment that is
bases is strongly dependent on the water content maintained by the presence of a high concentra-
of the considered microorganisms, with two tion of calcium dipicolinate (Ca-DPA). It was
extreme situations that involve vegetative cells also shown that Ca-DPA is able to photosensitize
and bacterial spores. the UVC-mediated formation of SP while
protecting DNA against the damaging effects of
Vegetative Cells UVB and UVA radiations (Cadet and Douki
Dimerization of two adjacent pyrimidine bases is 2010). It may be added that CPDs and 6-4PPs
the overwhelming reaction induced upon expo- are formed in significant amounts in spores that
sure of DNA of vegetative cells to UVC and UVB lack either SASP or Ca-DPA, showing the major
photons, thus giving rise to two main types of role played by the latter molecules in the gener-
photoproducts, namely cis-syn cyclobutane ation of SP.
pyrimidine dimers (CPDs) and pyrimidine (6-4)
pyrimidone photoproducts (6-4PPs) (Fig. 1). The Radiation-Induced DNA Damage
formation of CPDs involves a [2 + 2] cycloaddi- The molecular effects of ionizing radiation on
tion between the C5-C6 double bonds of the two cellular DNA may be rationalized in terms of
adjacent pyrimidine bases. The generation of indirect effects through the generation of
6-4PPs is rationalized in terms of [2 + 2] Patern- hydroxyl radical (•OH), the product of water mol-
ò-B€
uchi cycloaddition between the C5-C6 double ecule radiolysis (Fig. 1), and direct interaction
bond of the 50 -end pyrimidine to either the C4 with the genetic material that leads to ionization
carbonyl of a thymine or the imine group of a of the bases and the 2-deoxyribose moiety (von
cytosine in a suitable tautomeric form (Cadet and Sonntag 1987). An important aspect to be con-
Vigny 1990; Taylor 1994). The efficiency of sidered in the mode of action of ionizing radiation
bipyrimidine photoproduct formation and the rel- is the multiplicity of radical and excitation hits
ative yield of CPDs and 6-4PPs are strongly that follow the energy deposition (Goodhead
dependent on the primary DNA sequence. 1994). As a result, clustered damage, initially
Another striking feature is the efficient hydrolytic called “multiply damage,” consisting of several
deamination of cytosine residues in either CPDs lesions (modified bases, abasic sites, single- and
or at the 50 -end of 6-4PPs that leads to the forma- double-strand breaks), may be generated within
tion of mutagenic uracil derivatives. One may one or two helix turns. It may be added that the
add that UVA is able to induce CPDs through a complexity of the clustered DNA damage
mechanism that remains to be elucidated with an increases with the ▶ linear energy transfer
efficiency that is higher than the generation of (LET) value of ionizing radiation and heavy par-
8-oxo-7,8-dihydro-20 -deoxyguanosine ticles. This is particularly of concern for space
(8-oxodGuo) (Cadet et al. 2009a). ionizing radiation that consists of high-energy
670 DNA Damage
DNA Damage,
Fig. 1 Photo- and
O
radiation-induced damage UV photons
N
OH
in DNA: UVB-induced (190 < λ < 320 nm)
N
bipyrimidine (1) (6-4)
O
photoproducts upon Photoproduct
N
HN
photoexcitation and
radiation-induced base and
O
O
2-deoxyribose degradation
NH
products by hydroxyl
radical (•OH) and
N
Absorption/excitation
O
one-electron oxidation
(ionization) (2) Cyclobutane
dimer
N
O
HN
O
g
izin
(1) Indirect Ion
effect
n
iatio
Rad
(3) Clustered
damage
(2) Direct
ionization
protons and densely ionizing high-LET HZE par- (Fig. 2) have been shown to arise from
ticles ranging from helium to uranium, whereas one-electron oxidation and one-electron reduc-
terrestrial radiation is represented mostly by tion, respectively, of 8-hydroxy-7,8-
low-LET photons (X-, b-, or g-rays) (Durante dihydroguanyl radical, the initial •OH addition
and Cucinotta 2008). product to guanine (Cadet et al. 2009b). Similar
degradation products are generated by •OH addi-
•
OH and One-Electron Oxidation Products as tion to adenine, however, with an efficiency that
One-Hit Lesions is about tenfold lower than for guanine degrada-
A large body of information is available on tion products. One-electron oxidation of
the •OH and one-electron oxidation of the nucleobases in cells gives rise to the same pattern
purine and pyrimidine bases of DNA, thanks to of degradation products as those induced by •OH.
comprehensive and detailed studies on model However, one may note a different product dis-
compounds (Cadet et al. 2009b; Wagner and tribution since 8-oxodGuo is formed predomi-
Cadet 2010). Thus, the 4 cis and trans diastereo- nantly upon ionization of the bases as the result
mers of 5,6-dihydroxy-5,6-dihydrothymidine, of positive hole migration along the DNA chain
5-(hydroxymethyl)-20 -deoxyuridine, and with preferential trapping of the radical cation at
5-formyl-20 -deoxyuridine have been character- guanine sites (Genereux and Barton 2010;
ized as the main •OH-mediated degradation prod- Kanvah et al. 2010). It is well documented that
ucts of thymidine in cellular DNA (Cadet DNA single-strand breaks (SSBs) are generated
et al. 2008). In addition, 8-oxo-7,8-dihydro- by •OH-mediated hydrogen abstraction from the
20 -deoxyguanosine (8-oxodGuo) and 2-deoxyribose moiety (von Sonntag 2006; Dedon
2,6-diamino-4-hydroxy-5-formamidopyrimidine 2008). Ionization reactions are also likely to be
DNA Damage 671
8-oxodGuo
O
O H
N
HN
°+ + H2 O O HN
N
Oxidation
N − H+ O
−e− H2N N N°
HN H2N N N
dR OH
H dR
dGuo H2N N
N
O H
dR
Reduction O
D
N
Neutral reducing radical HN
HO°
H
H2N N NH
dR
FapydGuo
DNA Damage, Fig. 2 Radiation-induced formation 4-hydroxy-5-formamidopyrimidine (FapydGuo) as the

of 8-oxo-7,8-dihydro-20 -deoxyguanosine (8-oxodGuo) result of either •OH addition or electron abstraction
and N(2-deoxy-b-d-erythro-pentofuranosyl)-2,6-diamino-
involved in the formation of SSBs although there apurinic sites on the two complementary DNA
is still a lack of mechanistic information. strands within one or two DNA helix turns (Hada
and Georgakilas 2008). Attempts have been
Specific Radiation-Induced Clustered Damage made using a DNA repair-based assay to monitor
DNA double-strand breaks (DSBs) that consist of the formation of these lesions. However, it was
2 closely spaced and opposed single-strand found that the yield of clustered lesions, thus
breaks (SSBs) separated by less than 15 base measured, is slightly lower to those of single
pairs are the most representative radiation- oxidized bases in irradiated cells, a rather surpris-
induced DNA clustered damage. It has been ing finding. At this point it may be stressed that
predicted by Monte Carlo calculations that the further work is required to better assess the for-
frequency of complex DSBs that contain at least mation of clustered DNA modifications.
one additional nick/or modified base increases up
to 70 % for 2 MeV particles, while the proportion
was only 20 % for low-LET 4.5 keV electrons. Applications
Experimental support for the induction of a
higher density of DSBs with the increase in The measurement of CPDs, 6-4PPS, and SP pho-
LET of incident HZE particles was provided by toproducts has been made in the DNA of several
the observation of a larger frequency of fluores- microorganisms allowing detailed studies of
cent g-H2AX foci (i.e., 1 of the eukaryotic his- their repair. The yields of bipyrimidine photo-
tones that becomes phosphorylated on serine products vary according to the microorganisms
139 as a reaction on DNA DSBs). It has also due to the presence in some cases of
been shown that the DSBs generated with high- photoprotective compounds and differences in
LET radiation are more difficult to be repaired the membrane composition. One may also note
likely due to an increase in the complexity of the that the radiation-induced formation of 8-
clustered lesions. oxodGuo in cellular DNA was found to decrease
A second class of complex DNA damage con- with the increase in LET of the radiation. This is
sists in non-double-strand break clustered dam- indirectly indicative of the major deleterious role
age that may comprise at least one strand break played by complex clustered DNA damage in
together with one or several base lesions and/or ▶ radiation biology.
672 DNA Damage Correction
Future Directions Cadet J, Sage E, Douki T (2005) Ultraviolet radiation-

mediated damage to cellular DNA. Mutat Res
571:3–17
Efforts should be made to better ascertain the Cadet J, Douki T, Ravanat J-L (2008) Oxidatively gener-
effects of vacuum-UV light on the DNA of resis- ated damage to the guanine moiety of DNA: mecha-
tant microorganisms including bacterial spores nistic aspects and formation in cells. Acc Chem Res
when exposed to outer space conditions in terms 41:1075–1083
Cadet J, Douki T, Ravanat J-L, Di Mascio P (2009a)
of photoproduct distribution. Another major topic Sensitized formation of oxidatively generated damage
to be further investigated concerns the detection to cellular DNA by UVA radiation. Photochem
and quantification of radiation-induced clustered Photobiol Sci 8:903–911
damage whose identification remains a challeng- Cadet J, Douki T, Gasparutto D, Ravanat J-L, Wagner JR
(2009b) Chemical reactions of the radical cations of
ing issue due to the complexity of the analytical nucleobases in isolated and cellular DNA. Formation
problem thus addressed and the oneness of each of single-base lesions. In: Greenberg MM (ed) Radical
of the damage. and radical ion reactivity in nucleic acid chemistry.
Wiley, Hoboken, pp 69–97
Cadet J, Douki T, Ravanat J (2010) Oxidatively generated
damage to DNA. Free Radic Biol Med 4(49):9–23
See Also Dedon PC (2008) The chemical toxicology of 2-
deoxyribose oxidation in DNA. Chem Res Toxicol
21:206–219
▶ Aerobiology Durante M, Cucinotta FA (2008) Heavy ion carcinogene-
▶ Cosmic Rays in the Heliosphere sis and human space exploration. Nat Rev Cancer
▶ DNA 8:465–472
▶ DNA Repair Genereux JC, Barton JK (2010) Mechanisms for DNA
charge transport. Chem Rev 110:1642–1662
▶ HZE Particle Goodhead DT (1994) Initial events in the cellular effects
▶ Ionizing Radiation, Biological Effects of ionizing radiations: clustered damage in DNA. Int
▶ Linear Energy Transfer J Radiat Biol 65:7–17
▶ Mutation Hada M, Georgakilas AG (2008) Formation of clustered
DNA damage after high-LET irradiation: a review.
▶ Nucleic Acids J Radiat Res 49:203–210
▶ Nucleic Acid Base Kanvah S, Joseph J, Schuster GB, Barnett RN, Cleveland
▶ Photobiology CL, Landman U (2010) Oxidation of DNA: damage to
▶ Photochemistry nucleobases. Acc Chem Res 43:280–287
Nicholson WL, Munakata N, Horneck G, Melosh HJ,
▶ Radiation Biology Setlow P (2000) Resistance of Bacillus endospores to
▶ Solar Particle Events extreme terrestrial and extraterrestrial environments.
▶ Solar UV Radiation, Biological Effects Microbiol Mol Biol Rev 64:548–572
▶ Space Environment Taylor J-S (1994) Unraveling the molecular pathway from
sunlight to skin cancer. Acc Chem Res 27:76–82
▶ Survival von Sonntag C (1987) The chemical basis of radiation
▶ UV Climate biology. Taylor & Francis, London
▶ UV Radiation Dose von Sonntag C (2006) Free radical induced DNA damage
▶ UV Radiation, Biological Effects and its repair. A chemical perspective. Springer,
Heidelberg
Wagner JR, Cadet J (2010) Oxidation reactions of cyto-
sine DNA components by hydroxyl radical and
References and Further Reading one-electron oxidants in aerated aqueous solutions.
Acc Chem Res 43:564–571
Cadet T, Douki T (2010) Molecular effects of UV and
ionizing radiation on DNA. In: Gargaud M, Lopez-
Garcia P, Matin H (eds) Origins and evolution of life:
an astrobiological perspective. Cambridge University
Press, Cambridge, pp 359–374
Cadet J, Vigny P (1990) The photochemistry of nucleic
acids. In: Morrison H (ed) Bioorganic photochemistry:
DNA Damage Correction
photochemistry and the nucleic acids, vol 1. Wiley,
New York, pp 1–272 ▶ DNA Repair
DNA Repair 673
See Also
DNA Polymerase
▶ DNA
Juli Peretó ▶ Replication (Genetics)
Institut Cavanilles de Biodiversitat i Biologia ▶ Template
Spain
D
DNA Recombination
Synonyms
▶ Recombination
DNA- (or RNA-)dependent DNA polymerase;
DNA replicase; Klenow fragment
DNA Repair
Definition
Wayne L. Nicholson1,2 and Ralf Moeller3
DNA polymerase is an enzyme which catalyzes 1
Space Life Sciences Laboratory, University of
the addition of a deoxynucleoside triphos-
phate to the 30 end of a DNA strand using a 2
Space Life Sciences Laboratory, Kennedy Space
template. The systematic name of this enzymatic
Center, University of Florida, Gainesville, FL, USA
activity is deoxynucleoside-triphosphate:DNA 3
deoxynucleotidyl transferase and the catalyzed
reaction can be represented as
Deoxynucleoside triphosphate þ DNAn

Keywords
¼ diphosphate þ DNAnþ1
DNA damage; DNA, repair of damage
DNA-directed DNA polymerases (EC 2.7.7.7)
use DNA templates, whereas RNA-directed
DNA polymerases (EC 2.7.7.49), also known as Synonyms
reverse transcriptases, use RNA templates. These
enzymes cannot initiate a chain de novo and DNA damage correction
require a primer, which may be DNA or RNA.
Some DNA-directed DNA polymerases have
acquired proofreading mechanisms through a Definition
30 –50 exonuclease activity.
▶ DNA repair refers to a collection of processes
by which a cell identifies and corrects damage to
History the DNA molecule(s) that encode its genetic
information.
DNA polymerase I from Escherichia coli was
discovered by Arthur Kornberg in 1956
(awarded with the Nobel Prize in Physiology or History
Medicine in 1959). Howard Temin and David
Baltimore independently discovered viral reverse The discovery and early history of DNA repair
transcriptase in 1970 (both scientists shared the has been reviewed recently (Friedberg 2008 and
Nobel Prize in Physiology or Medicine in 1975). references therein).
674 DNA Repair
Overview • Excision repair pathways are used to correct a

number of different types of damage occurring
Deoxyribonucleic acid (DNA) is the on only one of the two DNA strands. In gen-
molecule that stores the genetic instructions eral, damage-specific enzymes excise the
used by all known living organisms and some damaged region and the correct sequence is
viruses (other viruses use ribonucleic restored by a specific repair DNA polymerase
acid – RNA – as their genetic material). Success- using the opposite, undamaged strand as a
ful propagation of life is therefore dependent template. Examples are ▶ nucleotide excision
upon the faithful copying of the information repair, base excision repair, and methyl-
encoded in the sequence of DNA bases in a pro- directed mismatch repair (Friedberg
cess called replication. et al. 2006).
In the context of astrobiology, DNA can be • DNA damage can also be corrected by homol-
damaged by exposure to environmental factors ogous ▶ recombination repair. In this path-
such as heat, ultraviolet or ionizing radiation, way, when DNA is being actively replicated,
extreme vacuum, desiccation, various toxic pieces of damaged DNA can be swapped for
chemicals, or oxidizing agents. DNA can also undamaged DNA originating from the
be damaged as a result of normal metabolic pro- undamaged daughter strand, using a series of
cesses occurring within cells. ▶ DNA damage specific recombination (Rec) enzymes
can either: (1) block the progress of replication (Friedberg et al. 2006).
by the ▶ enzyme DNA polymerase, leading to • Double strand breaks (DSB) can be formed in
cell death; or (2) lead to the incorporation of DNA by ionizing radiation or extreme desic-
incorrect bases during replication, resulting in cation. DSB are especially lethal because they
mutations. disrupt the integrity of the DNA molecule.
In order to maintain the integrity of the DSB are repaired in bacteria by a pathway
structure and base sequence in DNA, the cell called non-homologous end joining (NHEJ),
employs a number of enzymatic DNA repair in which the broken ends are rejoined by a
processes to specifically recognize and correct specific DNA ligase. NHEJ has been shown
damage to DNA or errors of base incorporation. to be important in the survival of spores to
Most of the DNA repair pathways described various types of ionizing radiation and high-
below have been shown to be important for energy particle bombardment (Moeller
microbial survival to exposure to outer space or et al. 2007).
other extreme environments such as the surface
of Mars (Nicholson et al. 2005; Moeller
et al. 2007):
See Also
• Certain types of DNA damage can be repaired
by direct reversal using enzymes that recog- ▶ DNA
nize the specific type of damage. Photoreacti- ▶ DNA Damage
vation is a light-dependent DNA repair ▶ Enzyme
mechanism, in which the UV-induced cis-syn ▶ Error Rate
cyclobutane pyrimidine dimers (CPDs) are ▶ Genetics
enzymatically monomerized by a photolyase ▶ Ionizing Radiation, Biological Effects
(Friedberg et al. 2006). UV-irradiated bacte- ▶ Mutagenesis
rial spores accumulate a spore-specific thy- ▶ Mutation
mine dimer, spore photoproduct (SP), that ▶ Nucleotide
can be directly reversed to two thymines ▶ Recombination
using the enzyme SP lyase (Nicholson ▶ Replication (Genetics)
et al. 2000). ▶ Solar UV Radiation, Biological Effects
Domain (Taxonomy) 675
References and Further Reading developed an alternative system which relies on

the abortive synthesis of complementary DNA
Friedberg EC (2008) A brief history of the DNA repair strains, thanks to the use of dideoxynucleotide
field. Cell Res 18:3–7
triphosphates – also called “dideoxys” – as
Friedberg EC, Walker GC, Siede W, Wood RD, Schultz
RA, Ellenberger T (2006) DNA repair and mutagene- chain terminators. In both methods, the fragments
sis, 2nd edn. ASM Press, Washington, DC are separated and identified by ▶ electrophoresis.
Moeller R, Stackebrandt E, Reitz G, Rettberg P, Doherty The Sanger method for DNA sequencing gave rise
AJ, Horneck G, Nicholson WL (2007) Role of DNA
to automatic DNA sequencers based on capillary
repair by non-homologous end joining in Bacillus D
subtilis spore resistance to extreme dryness, mono- electrophoresis, thus triggering the explosion of
and polychromatic UV and ionizing radiation. ▶ genome sequencing projects in the 1990s.
J Bacteriol 189:3306–3311 Since 2005, a number of “next-generation” tech-
Nicholson WL, Munakata N, Horneck G, Melosh HJ,
niques have been released, producing a spectacu-
Setlow P (2000) Resistance of Bacillus endospores to
extreme terrestrial and extraterrestrial environments. lar increase in sequencing speed together with a
Microbiol Mol Biol Rev 64:548–572 drastic reduction in the cost. They include the
Nicholson WL, Schuerger AC, Setlow P (2005) The solar so-called pyrosequencing, sequencing by synthe-
UV environment and bacterial spore UV resistance: con-
sis, and sequencing by ligation. Later on, the cur-
siderations for Earth-to-Mars transport by natural pro-
cesses and human spaceflight. Mutat Res 571:249–264 rent “third-generation” techniques allow the fast
and reliable sequencing of single DNA molecules.
This is widely regarded as the future of sequenc-
ing, since single-molecule approaches do not
DNA Replicase require the DNA to be previously amplified.
▶ DNA Polymerase
See Also
▶ Amplification (Genetics)
DNA Sequencing
▶ DNA
▶ Electrophoresis
Carlos Briones
▶ Genetics
Centro de Astrobiologı́a (CSIC/INTA), Consejo
▶ Genome
Superior de Investigaciones Cientı́ficas, Madrid,
▶ Mutation
Spain
▶ Nucleotide
▶ Sequence
Synonyms
Gene sequencing; Sequencing

Domain (Taxonomy)
Definition Purificación López-Garcı́a

Unité d’Ecologie, Systématique et Evolution,
▶ DNA sequencing is the process by which the CNRS UMR8079 Université Paris-Sud 11, Paris,
order of ▶ nucleotides in a DNA molecule is Orsay Cedex, France
determined. The first DNA sequencing
method – set up by A. Maxam and W. Gilbert in
the decade of the 1970s – was based on the Keywords
controlled chemical breakage at one or two of
the four ▶ nucleotides. In parallel, F. Sanger Archaea; Bacteria; Eukarya
676 Doppler Shift
Synonyms (Archaea and Bacteria) including cell wall bio-

chemistry and the presence of eukaryotic-like
Empire; Primary kingdom; Superkingdom RNA polymerases in Archaea. The marked differ-
ences between archaea and bacteria have been
widely demonstrated since then, thanks, among
Definition others, to the comparison of full genome
sequences. The classification of life in three phylo-
Domain is the highest taxonomic rank in the genetic domains is well established today. It does
hierarchical biological classification system, not imply the demise of the prokaryote-eukaryote
above the kingdom level. There are three dichotomy at the structural level, which is an
domains of life, the ▶ Archaea, the ▶ Bacteria, empirical reality. Therefore, there are two types
and the ▶ Eukarya. Organisms from Archaea and of cells from a structural point of view, but three
Bacteria have a prokaryotic ▶ cell structure, domains of life from a phylogenetic point of view.
whereas organisms from the domain Eukarya
(eukaryotes) encompass cells with a nucleus con-
fining the genetic material from the cytoplasm. See Also
▶ Archaea
Overview ▶ Bacteria
▶ Cell
The term “domain” was introduced by Carl ▶ Eukarya
R. Woese et al. (1990) together with the proposal ▶ Phylogeny
of a natural classification system for all life on ▶ Taxonomy
Earth, including microorganisms, which had previ-
ously escaped any attempt of classification based
on evolutionary relationships (Woese et al. 1990).
Woese’s proposal to class life in three major phy- Sapp J (2009) The new foundations of evolution. Oxford
logenetic domains was an attempt to institutional- University Press, New York, p 425
ize the three major phylogenetic groupings of Woese C, Fox GE (1977) Phylogenetic structure of the
organisms that he had previously observed and prokaryotic domain: the primary kingdoms. Proc Natl
Acad Sci U S A 74:83–86
defined (Woese and Fox 1977) based on differ- Woese CR, Kandler O, Wheelis ML (1990) Towards a
ences in the small subunit ribosomal RNA, a mac- natural system of organisms. Proposal for the domains
romolecule chosen as an ideal phylogenetic marker Archaea, Bacteria and Eucarya. Proc Natl Acad Sci
due to its universality, degree of conservation, and U S A 87:4576–4579
essential function. The three-domain classification
system was proposed as an alternative to other life
classification systems in use at that time, such as
the traditional prokaryote-eukaryote division or the Doppler Shift
five-kingdom system (Monera, Protista, Fungi,
Plantae, and Animalia) proposed by R. Whittaker Daniel Rouan
in 1969. The three-domain classification based on LESIA, Observatoire Paris-Site de Meudon,
ribosomal RNAs was received with skepticism by Meudon, France
most biologists at that time, who were not ready to
accept that the evolutionary information stored in
one single macromolecule had more weight than Definition
other, phenotypic, characters. O. Kandler and his
German school provided additional features that The Doppler shift is the change in frequency and
distinguished radically the two prokaryotic groups wavelength of a propagating wave (acoustic,
Dormant State 677
Doppler Shift,
Fig. 1 Diagram
illustrating how the
measured wavelength of a
wave emitted by a moving
source changes, depending
on the direction of the
movement with respect to
the observer
D
electromagnetic, etc.), caused by the movement See Also

of the source emitting the wave relative to
the observer. This effect is named after the ▶ Radial Velocity
Austrian physicist Christian Doppler who ▶ Radial-Velocity Planets
proposed it in 1842. In France, the effect is cur-
rently called Doppler–Fizeau, because the
French physicist Hippolyte Fizeau proposed it
for electromagnetic waves in 1848. If the
source is moving away from the observer, the
wavelength appears stretched while frequencies Dormant State
are shifted toward lower values, and if the
source is approaching, the wavelength Felipe Gomez
appears compressed and frequencies shifted Centro de Astrobiologı́a (CSIC/INTA), Instituto
toward higher values. In the case of light, the Nacional de Técnica Aeroespacial, Torrejón de
effect is called redshift (receding source) and Ardoz, Madrid, Spain
blueshift (approaching source), respectively.
This effect is of great importance in
astronomy because it allows direct determination Definition
of the radial speed of celestial objects and of the
distance of remote galaxies (because of the Dormant state refers to a reduced metabolic
expansion of the Universe). It provides activity phase adopted as strategy to escape
the basis for the most productive method for stressful conditions and/or save energy. It refers
detection of exoplanets, the so-called ▶ radial to a latent but capable to be active state. The
velocity method. The relative amount of wave- adverse weather conditions (low temperature in
length change when the velocity is not close to winter time, etc.. . .) drive a plant or animal pop-
the speed of light is given by the relation Dl/l = ulation to reduce their metabolic activity in order
v/c, where v is the relative velocity, c is the to preserve themselves avoiding frosting condi-
velocity of the wave (e.g., speed of light), l is tions. The spore state of sporulating Gram-
the wavelength, and Dl, the change in wave- positive bacteria is considered a dormant state
length (Fig. 1). of the bacterium.
678 Double Helix
the B-form, while double-stranded ▶ RNA typi-

Double Helix cally adopts the A-form.

Earth–Life Science Institute (ELSI), Tokyo See Also
Japan ▶ Base Pair
Institute for Advanced Study, Princeton, NJ, ▶ DNA
USA ▶ Nucleic Acids
Blue Marble Space Institute of Science, ▶ Watson-Crick Pairing
Washington, DC, USA
Drake Equation
Definition
Leticia Carigi
Instituto de Astronomı́a, Universidad Nacional
A double helix consists of two congruent helices
Autónoma de México, México, DF, Mexico
with the same axis, differing by their translation
along the axis. In molecular biology, complemen-
tary double-stranded ▶ DNA typically adopts a
Keywords
double helical structure. The DNA double helix
can be (from left to right in Fig. 1 below) of the
Extraterrestrial civilization
A- or B-form, which are right-handed but differ
from each other in the distance required to make a
complete helical turn or of the Z-form, which is
Synonyms
left-handed. The double helix model of DNA
structure was published in Nature by Watson
Green bank equation
and Crick in 1953. DNA is typically found in
Double Helix,
Fig. 1 Three double helix
structures adopted by
double stranded DNA
under different
physiological conditions
DTU Space, Denmark 679
Definition the habitable planets or moons in stars unlike our

Sun, the change of the habitable zone with the
The Drake equation estimates the number of evolution of the star, the metallicity requirements
communicating extraterrestrial civilizations that for the formation of earthlike planets, etc.
might exist within the ▶ Milky Way today. That
formula was devised by Frank Drake in 1961.
See Also
Overview ▶ Exoplanets, Discovery D

▶ Galactic Habitable Zone
The original Drake equation gives the number ▶ Habitable Zone
(N) of civilizations in the Galaxy that are able to ▶ Search for Extraterrestrial Intelligence
communicate through electromagnetic waves. That ▶ SETI
formula is written as the product of seven terms, as
follows: N ¼ R f p ne f l f i f c L ,
where (1) R is the rate of formation of stars suitable
for life, that is, solar-type stars, in units of stars per Drake FD (1962) Intelligent life in space. Macmillan, New
year; (2) fp is the fraction of suitable stars York
surrounded by planetary systems; (3) ne is the num- Konesky G (2009) The Drake equation revisited. SPIE
ber of planets suitable for life in a given planetary Proceedings, 7441
Lemarchand G (2004) The technological adolescent age
system, that is, the number of planets located within transition: a boundary to estimate the last factor of the
the ▶ habitable zone; (4) fl is the fraction of habit- Drake equation. IAUS 213:460–466
able planets, within that zone, where basic life
appears; (5) fi is the fraction of planets on which
basic life evolves to intelligent life; (6) fc is the
fraction of planets on which intelligent life develops DTU Space, Denmark
a sufficiently high technology capability to broad-
cast signals of its existence out into the Galaxy; and Michel Viso
(7) L is the lifetime of such a technological civili- CNES/DSP/SME, Vétérinaire/DVM, Astro/
zation emitting detectable signals into space. Exobiology, Paris Cedex 1, France
The R factor is the best understood due to the
vast body of astronomical data available; fp and
ne factors can be estimated with some degree of Synonyms
certainty using the properties of exoplanets
detected so far, However, the remaining factors Institut for Rumforskning og -teknologi på
are unknown at present since we do not know any Danmarks Tekniske Universitet
extra terrestrial life form or civilization.
Thus, depending on whether one adopts the
pessimistic or the optimistic point of view, a Definition
civilization may last 10 years or 100,000 years;
he/she may obtain N = 1 or N = 10,000, respec- Until 2007, the Danish National Space Center
tively, indicating that we are either almost alone (DNSC) was an independent government research
in the Milky Way or, on the contrary, in the institute with its roots dating back to the beginning
Galaxy there exist an abundance of extraterres- of the space age in 1962. These roots include
trial civilizations that have yet to be detected. Danish Space Research Institute (DSRI) from
An important problem with the Drake equation is 1968 to 2005, which, by merging in 2005 with
its incompleteness since the formula omits impor- two research units of the Danish National Survey
tant factors such as possible interstellar colonization, and Cadastre, and by adding the fields Geodesy
680 Dual Status Objects
and Geodynamics, created the DNSC. On January

1, 2007, parts of Ørsted•DTU and the Department Dust Cloud, Interstellar
of Informatics and Mathematical Modeling at
DTU merged with the Danish National Space William M. Irvine
Center. The recent mergers have brought together University of Massachusetts, Amherst, MA, USA
most space related activities in Denmark, now
forming the National Space Institute (NSI) with a
staff of more than 110 persons.
Synonyms
Dark cloud
Dual Status Objects
▶ Active Asteroid Definition
Interstellar molecular clouds which lack obvious

evidence of recent star formation, and hence
Dubiofossil which are observable because their dust grains
block the light from background stars, are some-
Emmanuelle J. Javaux times called dust clouds. Note that all ▶ molecu-
Palaeobiogeology-Palaeobotany- lar clouds do contain dust grains, whether or not
Palaeopalynology, Geology Department, they also contain stars.
See Also
Definition
A dubiofossil is a problematic structure that looks ▶ Interstellar Medium
like a ▶ fossil but whose biological origin is ▶ Molecular Cloud
uncertain or ambiguous. Several physical and
chemical processes may produce structures
resembling biological structures. Detailed ana-
lyses and criteria of ▶ biogenicity have to be
Dust Devils
applied on the dubiofossils to move them to the
category of fossils (biogenic or biological struc-
Barbara Stracke
tures) or ▶ pseudofossils (abiogenic or
nonbiological structures). This is a crucial point
when looking for early traces of life on Earth or
Germany
for extraterrestrial life.
See Also Definition
▶ Biogenicity Dust Devils are meteorological phenomena that

▶ Biomarkers have been detected in the atmospheres of ▶ Earth
▶ Fossil and ▶ Mars. They are thermally driven atmo-
▶ Microfossils spheric vortices that are filled with loose mate-
▶ Pseudofossil rials such as dust and sand. The vertical velocity
Dwarf Planet 681
is predominantly upward. Their size ranges

with heights from a few meters (on Earth and D-Value
Mars) to several kilometers (only on Mars) and
from a few meters to hundreds of meters in diam- Catharine A. Conley
eter. The first images of Dust Devils in the NASA Headquarters, Washington, DC, USA
Martian atmosphere were taken by the ▶ Viking
orbiter.
Definition
D
The D-value is a number given to characterize a
See Also
process to achieve inactivation of 90 % of a
population of the test ▶ microorganism under
▶ Earth
stated conditions (temperature, pressure, etc.).
▶ Mars
This value could be a dose or a concentration
▶ Viking
for a drug, a time in minute, hour, or day for
▶ DHMR, or pasteurization, a dose for ▶ sterili-
zation using ionizing radiations (i.e., g-ray).
Dust Grain
See Also
William M. Irvine
▶ Bioburden
▶ DHMR
▶ Microorganism
Synonyms
▶ Sterilization
Interstellar grain
Dwarf Planet
Definition
Tilman Spohn
The small solid particles present in the interstellar
medium are called dust grains or often simply
either dust or grains. Most of them are only a
Germany
few tenths of a mm in size. They consist of sili-
cates and carbon compounds, coated in the
coldest regions with icy mantles. They represent
Definition
1–2 % of the mass of interstellar clouds. Their
origin and composition are discussed in interstel-
A dwarf planet according to the IAU
lar dust.
(International Astronomical Union) Resolutions
5 and 6 (Resolution_GA26-5-6, passed in 2006)
is a celestial body of intermediate size, which
See Also orbits the Sun, is not a satellite, has not cleared
the neighborhood around its orbit, and has suffi-
▶ Interstellar Dust cient mass for its self-gravity to maintain a nearly
▶ Interstellar Ices round shape. To date five dwarf planets have been
682 Dwarf Star
identified, namely, the asteroid ▶ Ceres, ▶ Pluto, See Also

Eris, Makemake, and Haumea, the latter three
being ▶ trans-Neptunian objects. ▶ Hertzsprung-Russell Diagram
▶ Main Sequence, Star
▶ Star
See Also ▶ Stellar Evolution
▶ Ceres
▶ Pluto
▶ Sun (and Young Sun)
▶ Trans-Neptunian Object Dynamical Friction
Sean N. Raymond
Laboratoire d’Astrophysique de Bordeaix,
Dwarf Star CNRS, Universite de Bordeaux, France
David W. Latham
Harvard-Smithsonian Center for Astrophysics, Definition
Cambridge, MA, USA
Dynamical friction is a force exerted by
small objects that damps the random motions of
Definition large objects. This occurs via a gravitational
“focusing” of small bodies behind the orbit of
Stars of a given mass are smallest in size when in the large body. This is different than
their main-sequence stage and are hence called viscous stirring, which is simply an equipartition
dwarfs; this use of the term should not be con- of energy between large and small bodies.
fused with ▶ white dwarfs, the end stage (death) In the context of planet formation, dynamical
of low-mass stars. Stars similar to the Sun spend friction from planetesimals acting on ▶ planetary
most of their luminous careers burning hydrogen embryos keeps the eccentricities and
in their cores. It is during this main-sequence or inclinations of the embryos relatively small
dwarf-star phase that the energy output can be such that collisions between embryos only occur
stable over periods of time long enough to sup- once the planetesimal population is largely
port the evolution of life as we know it. Dwarf depleted.
stars are prime targets for searches for
▶ transiting planets as small as the Earth. Recent
attention has focused on the coolest dwarf stars of
spectral type M, because they range from one half
See Also
to one tenth the diameter of the Sun. Planetary
transits around such stars are thus much deeper ▶ Planetesimals
▶ Viscous Stirring
and easier to detect; in addition, for a given plan-
etary mass, the wobble of the star will be larger,
making radial velocity and astrometry measure-
ments easier. The ▶ habitable zone where water
could be liquid is also much closer to a cool star,
with a correspondingly shorter orbital period and Goldreich P et al (2002) Formation of Kuiper-belt binaries
higher likelihood to be correctly inclined to show by dynamical friction and three-body encounters.
transits. Nature 420:643–646
Dynamo, Planetary 683
Synonyms
Dynamical Instability
Magnetic field generation
Rory Barnes
Washington, Seattle, WA, USA Definition
A dynamo – generally – generates electric and

Definition magnetic energy from kinetic energy through D
the physical process of electromagnetic induc-
A dynamical instability is a nonlinear response to tion. A current will be induced in a closed wire
a phenomenon that ultimately leads to a large- moving through a magnetic field. The current by
scale change in a system. This term may be itself will generate a magnetic field. In planetary
applied to any type of dynamical system, such interiors, a similar although more involved
as orbits, atmospheres, interiors, etc. For exam- mechanism can produce a global ▶ magnetic
ple, during planet formation, the dust layer of a field; convective flow of an electrically
▶ protoplanetary disk is compressed, increasing conducting material in a magnetic field will
the density and ultimately coalescing the dust induce a current that will generate a magnetic
into ▶ planetesimals. On a large scale, if the gas field. If certain conditions are met, the magnetic
layer becomes cold enough and is massive field will maintain or amplify the original mag-
enough, the gas can collapse into a ▶ giant planet. netic field.
After planet formation if two planets come close
to each other, their orbits can be dramatically
altered, perhaps even resulting in one being Overview
ejected from the system. In these examples, the
particles (dust, gas, or planet) had been orbiting Generation of a magnetic field requires an elec-
the star in the same way for a long time, but the trically conducting shell within a ▶ planet,
orbit, or the particle itself, was quickly (usually in motion within that shell, and a “seed” magnetic
just a few to a few thousand orbits) changed. field. For recent reviews on dynamo problems,
see Busse and Simitiev (2015), Christensen and
Wicht (2015), and Roberts (2015). In the ▶ ter-
See Also restrial planets and the satellites, this region is
agreed to be the fluid iron-rich core at the center.
▶ Ejection, Hyperbolic In the ▶ Giant Planets ▶ Jupiter and ▶ Saturn, the
dynamo region is agreed to be the metallic
▶ hydrogen shell and in ▶ Uranus and ▶ Neptune
a comparatively shallow layer of ionized fluid
Dynamo, Planetary (see, e.g., Connerney 2015 for a review). In the
terrestrial planets and satellites, there may be a
Tilman Spohn solid inner core, the growth of which would pro-
Deutsches Zentrum f€ur Luft- und Raumfahrt vide a buoyancy flux that may drive the dynamo.
ur Planetenforschung, Berlin, This buoyancy then derives from a difference in
Germany composition between the solid inner core and the
fluid outer core. Light alloying elements such as
sulfur and oxygen tend to be expelled from the
Keywords solidifying core and to form a buoyant layer
above the inner core that tends to rise and mix
Planetary magnetic field; Planetary magnetism with the outer core. In addition to (or instead of) a
684 Dynamo, Planetary
chemical buoyancy flux from the inner core, ther- supplemented by conservation of mass and
mal buoyancy may drive the flow. The thermal entropy equations and by an equation of state.
buoyancy would result from a sufficiently large The magnetic induction equation (written in a
temperature difference between the core and the simplified form assuming a solenoidal velocity
silicate rock ▶ mantle surrounding the core. Fluid field and a constant magnetic diffusivity l) is
cooled at the core-mantle boundary will tend to
sink toward the center and thereby drive the con- @B
¼ ∇ ðv BÞ þ l∇2 B: (2)
vective flow. Thus, an entirely liquid core can @t
support a thermal dynamo. There is a difference
in efficiency between the thermal dynamo and the A sustained magnetic field can be generated by
compositionally driven dynamo. While the ther- the flow if the production of field energy by
mal dynamo as a heat engine is subject to a shearing the field (first term on the right-hand
Carnot efficiency (about 0.05, Gubbins side) overcomes the magnetic diffusion (second
et al. 2003), a compositionally driven dynamo is term on the right-hand side). The first term also
not as restricted. In the giant planets, thermal shows that certain restrictions apply to the flow.
buoyancy is usually thought to drive the flow A flow directed along the field lines, for instance,
although helium droplets falling out of solution will not contribute to the dynamo. The relative
with hydrogen may provide a buoyancy flux (see weight of the two terms is measured by the mag-
Guillot and Gautier 2015 for a review). It should netic Reynolds number
be mentioned that precession and tides have also
been suggested to excite flow instabilities in the Ud
Re ; (3)
core and a dynamo (e.g., Tilgner 2015 for a l
review).
Convection in the dynamo region can be where U is a characteristic velocity and d a char-
described by the field equations of fluid dynamics acteristic length (e.g., the core radius). The mag-
(e.g., Busse and Simitiev 2015). Because of the netic Reynolds number must be larger than 1 for
magnetic field, the electric currents, and the plan- the dynamo to operate. Other dimensionless
etary rotation, the Lorentz and Coriolis force parameter groups of relevance to the dynamo
have to be included. The momentum equation problem are the Rayleigh number that measures
then is the vigor of the convection and the Ekman num-
ber that measures the importance of the Coriolis
force.
@
r þ v ∇ v þ 2rV v ¼ ∇p þ rg þ j B þ rF: Numerical models of the dynamo driven by a
@t
buoyancy flux from below (a growing inner core)
(1)
have been successful, but the models are still far
In Eq. 1, t denotes time, v is the fluid velocity, r is from realistic values of both the Ekman and Ray-
density, V is the planet’s rotational angular leigh numbers. Nevertheless, a scaling law has
velocity vector, p is pressure, g is gravity, j is been derived for the magnetic field strength at the
the electric current density, B is the magnetic surface of a core dynamo region (Christensen and
induction vector, and F is the specific body Aubert 2006):
force (buoyancy). Vector quantities are written 13
in bold face. The first term on the left-hand side is 1 1 gQB d
B 0:9m r2 6 ; (4)
the inertia term, and the second term denotes the 4pRc Ri
Coriolis force. The first two terms on the right-
hand side denote pressure variations and where QB is the buoyancy flux into the dynamo
the hydrostatic pressure. The third term is the region (thermal and/or chemical) and Rc and Ri
Lorentz force, where mj ¼ ∇ B , with m the are the outer and inner radii of the core shell. The
magnetic permeability. The equation is characteristic length d here is the thickness of the
Dynamo, Planetary 685
fluid core shell RcRi. In this scaling, the mag- ▶ Heat Flow, Planetary
netic field strength depends on the size of the core ▶ Heat Transfer, Planetary
and the buoyancy flux but is independent of the ▶ Hydrogen
electrical conductivity and of the rotation rate, ▶ Jupiter
although it is still required that the magnetic ▶ Magnetic Field, Planetary
Reynolds number be larger than 1 and that in ▶ Mantle
order to obtain a dipolar field, the Coriolis force ▶ Neptune
must dominate over inertia. (The latter require- ▶ Planet
ment is consistent with Cowling’s theorem; see ▶ Rotation Planet D
Roberts 2015.) The scaling is similar in concep- ▶ Satellite or Moon
tion to a scaling introduced by Stevenson ▶ Saturn
et al. (1983), although the exponents in the ▶ Terrestrial Planet
power law differ. It is important to note that this ▶ Tides, Planetary
scaling supersedes an older scaling law that was ▶ Uranus
derived from a balance between the Lorentz force
and the Coriolis force terms in Eq. 1. In the latter
scaling – often used in the astrobiology literature
when discussing the habitability of Busse FH, Simitiev R (2015) Planetary dynamos. In:
exoplanets – the magnetic field was considered Schubert G, Spohn T (eds) Treatise on geophysics:
proportional to the planetary rotation rate (see planets and moons, vol 10. Elsevier, Amsterdam,
Hubbard and McFarlane 1980). pp 239–254
Christensen UR, Aubert J (2006) Scaling properties of
As Eq. 4 shows, the magnetic field strength convection-driven dynamos in rotating spherical shells
depends on the rate of heat transfer through the and application to planetary magnetic fields. Geophys
mantle of a terrestrial planet or satellite. The J Int 166(1):97–114
buoyancy flux is directly related to the ▶ heat Christensen UR, Wicht J (2015) Numerical dynamo sim-
ulations. In: Schubert G, Olson P (eds) Treatise on
flow extracted by the planetary mantle from the geophysics: core dynamics, vol 8. Elsevier, Amster-
core for a thermally driven dynamo. For a chem- dam, pp 245–277
ically driven dynamo, the rate of inner core Connerney JEP (2015) Planetary magnetism. In:
growth and therefore the rate of buoyancy release Schubert G, Spohn T (eds) Treatise on geophysics:
planets and moons, vol 10. Elsevier, Amsterdam,
depend on the core cooling rate and pp 195–237
therefore – again – on the heat flow from the Gubbins D, AlfË D, Masters G, Price D, Gillan MJ
core. The heat flow from the core can be calcu- (2003) Can the earth’s dynamo run on heat alone?
lated from thermal history models. This under- Geophys J Int 155:609–622
Guillot T, Gautier D (2015) Giant planets. In: Schubert G,
lines an important property of terrestrial planets: Spohn T (eds) Treatise on geophysics: planets and
A terrestrial planet may or may not have a mag- moons, vol 10. Elsevier, Amsterdam, pp 529–557
netic field, and the existence of a field is not easily Hubbard WB, McFarlane JJ (1980) Structure and
predicted from observations of bulk planetary evolution of uranus and neptune. J Geophys Res
85:225–234
parameters. Roberts PH (2015) Theory of the geodynamo. In:
Schubert G, Olson P (eds) Treatise on geophysics:
core dynamics, vol 8. Elsevier, Amsterdam, pp 57–90
See Also Stevenson et al (1983) Magnetism and thermal evolution
of the terrestrial planets. Icarus 54(3):466–489
Tilgner A (2007) Rotational dynamics of the core. In:
▶ Core, Planetary Schubert G, Olson P (eds) Treatise on geophysics:
▶ Exoplanets, Discovery core dynamics, vol 8. Elsevier, Amsterdam,
▶ Giant Planets pp 183–212
E
E there are numerous centers of excellence in astro-

biology in Europe, founders decided to promote
▶ Redox Potential their work by setting up this European Network to
help the sharing of their expertise and facilities.
This network aims also to attract young scientists
to participate practically in this evolving interdis-
ciplinary field. Other goals are the promotion of
Ea astrobiology to European funding agencies (ESA,
ESF) and political bodies such as the European
▶ Activation Energy Commission. Finally, individual members as
national associations are invited to promote a pub-
lic interest in astrobiology.
Up to 2010, scientists or organizations of
19 European countries are participating in
EANA this network: Austria, Belgium, Czech Republic,
Denmark, Finland, France, Germany, Greece,
Michel Viso Hungary, Italy, Portugal, Spain, Sweden,
CNES/DSP/SME, Vétérinaire/DVM, Switzerland, The Netherlands, Poland, Romania,
Astro/Exobiology, Paris Cedex 1, France Russia, and the UK. EANA holds annual work-
shops alternately in the different member countries.
EANA was established as an affiliate partner
Definition of the ▶ NASA Astrobiology Institute in early
2002. EANA is also associated with the Commis-
The European Astrobiology Network Association sion of Astrobiology, Space Life Origin and Evo-
(EANA) was organized in Spring 2001 during lution at the Chinese Society of Space Research,
the First European Workshop on Astrobiology and the Japanese Astrobiology Network.
co-organized with ▶ ESA at the ESRIN research For further information, visit http://www.
facility in Frascati, Italy. Acknowledging that astrobiologia.pl/eana/.

DOI 10.1007/978-3-662-44185-5
688 Early Archean
Early Archean Earth, Age of
▶ Barberton Greenstone Belt, Traces of Early Francis Albarède

Life Ecole Normale Supérieure de Lyon, Lyon,
France
Early Earth Keywords

▶ Archean Tectonics Accretion (planetary); CAIs; Chondrules; Decay
▶ Cool Early Earth constant; Geochronology; Meteorite; Radioactiv-
▶ Hadean ity; Radiogenic isotopes
Early Life Definition
▶ Barberton Greenstone Belt, Traces of Early The age at which material condensed from the
Life ▶ Solar Nebula ceased to significantly add to the
Earth.
Earth History
Nicholas Arndt Around 1850, British physicist Lord Kelvin

ISTerre, Université Grenoble Alpes, France (William Thomson) elaborated on the theory
of heat conduction formulated by the French
physicist and mathematician Joseph Fourier
Definition and concluded that the Earth was no more than
a few tens of Ma old. Kelvin also fiercely
The Earth is the planet on which we live. It is the opposed the geologists who guessed from mod-
third planet from the Sun and one of the telluric ern sedimentation rates that the Earth was older
(small and rocky) planets of our ▶ Solar System. than a billion years. However, by World War II,
It has a mean radius of 6,371 km and a mean the general agreement among scientists was that
density of 5.515 g cm3. It is distinguished from the Earth was older than 3 billion years. It is
other planets by the presence of ▶ continental often incorrectly stated that Kelvin’s mistake,
crust, oceans, and ▶ life. Its unusually large core one of the most devastating in the history of
and satellite (the Moon) are believed to result modern physics, was due to his ignorance of
from the collision of a Mars-sized body radioactive heating. It was British geologist
(protoplanet ▶ Theia) about 4.45 Ga ago. Arthur Holmes who pointed out that the loss of
heat from the interior of the Earth is largely
See Also controlled by mantle convection. In 1954,
American geochemist Clair Patterson brought
▶ Continental Crust the controversy to an end by showing that mete-
▶ Moon, The orites and the Solar System are some 4.55 Ga
▶ Terrestrial Planet old and that the age of the Earth cannot be very
▶ Theia different.
Earth, Formation and Early Evolution 689
Overview See Also
Even though the age of our planet is occasion- ▶ Earth’s Atmosphere, History of the Origins
ally presented as controversial, mostly on reli- ▶ Geochronology
gious grounds, all scientists agree that it started ▶ Radioactivity
forming some 4,567 million years ago. Two
real issues cloud the discussion. First, there is
no original rock sample of that age left on our
planet: the oldest crustal segment is 3.8 Ga old Dalrymple GB (1994) The age of the Earth. Stanford
(Isua Supracrustal Belt, West Greenland), the University Press, Stanford
Richter FM (1986) Kelvin and the age of the Earth. J Geol
E
oldest rock is some 4.1 Ga old (Acasta gneiss,
Canada), while the oldest undisputable mineral 94:395–401
ages are 4.3 Ga old detrital zircons (Jack Hills,
Western Australia). The problem of the age of
the Earth is therefore best addressed by dating
the earliest objects of the Solar System, the Earth, Formation and Early Evolution
calcium-aluminum-rich refractory inclusions
(▶ CAIs) found in some chondritic meteorites Stephen Mojzsis
and applying a comparative chronology based University of Colorado, Boulder, CO, USA
on extinct radionuclides to both terrestrial sam-
ples and meteorites in order to date the Earth’s
formation. The U-Pb age of CAIs is rather well Keywords
established at 4,568.0 0.5 Ma. The small
glass blebs found in chondrites, which are Acasta; Akilia; Basalt; Continents; Core
known as ▶ chondrules, are 1–3 Ma younger (planetary); Cosmochemistry; Crust; Early
than the CAIs. The ▶ radioactivity of the Earth; Earth’s atmosphere (origin and evolution
extinct 182Hf nuclide (T 1/2 = 8.5 Ma) requires of); Geochronology; Granite; Hadean; Impact
that the terrestrial core segregated 30 Ma after processes; Isotopes; Isua; Jack Hills; Late heavy
the formation of CAIs, which places a strong bombardment; Moon-forming impact; Oceans
constraint on the age of the planet itself. (origin of); Solar System; Zircon; Biopoesis;
Similar, but less tight constraints are placed Geochemistry; Lithosphere; Protolith
by other extinct nuclides (129I, 244Pu) and by
235
U. The early Solar System evolved by con-
densation of the nebular gas followed by dust Definition
agglomeration and then by coalescence of
asteroid-sized bodies and their merging into a Earth formed as a silicate- and metal-rich body
few tens of protoplanets. A second issue therein the context of the other inner solar system
fore is a matter of definition: how long did the “terrestrial” worlds. In its early evolution, it
accretion stage last? Astronomical evidence separated into layers (core, ▶ mantle, ▶ crust)
indicates that UV and X-rays emitted by as a consequence of the chemical and mechan-
proto-(T-Tauri)-stars similar to the Sun had ical properties of the materials that accreted
blown off the nebular gas and that the disk of to the Earth. The surface zone stabilized within
debris was swept clean in less than 5 Ma. the first 200 million years of Earth’s exis-
Dynamic models suggest that the period during tence and has hosted geochemical cycles mod-
which protoplanets merged to form planets as ulated in whole or in part by the presence of
we know them today may have lasted for a few liquid water and an atmosphere since about
extra tens of Ma. 4.4 Ga ago.
690 Earth, Formation and Early Evolution
Overview impacts which mitigated its long-term survival

which may be the reason why the oldest surviving
Introduction terrestrial rocks or minerals date from 150 Ma
Earth is a natural and accessible planetary-scale after accretion (Harrison 2009). The long resi-
laboratory to test ideas to be used in the search for dence times of the oldest rocks in the crust
life elsewhere in the Solar System and beyond. means that almost any geological process could
An understanding of Earth’s origin is essential to have been experienced by them. Over the inter-
the discipline of astrobiology, at least for the vening several billion years, ▶ weathering, denu-
reason that this planet is the singular known dation, erosion, ▶ metamorphism (thermal,
repository of life. Studies of the origin and early chemical, pressure induced, or all of the above),
evolution of Earth have undergone dramatic plate tectonics’ subduction, remelting, and other
shifts in recent years and have radically changed crustal recycling processes had been at work to
our knowledge of its formative time and how consume all but a minute fraction of what once
early events led to a habitable planet. This knowl- existed from the first billion years of geologic
edge logically forms the basis of ongoing time. Investigations focused on the origin of the
searches for other habitable solar system’s hydrosphere and on when the crust evolved into
exoplanets in the galactic neighborhood. More the current basaltic (oceanic) and granitic
than 400 planets around other stars have been (continental) dichotomy rely almost completely
documented by various methods (Schneider on information gleaned from a few outcrops or
2010) mostly within a sphere of space around isolated minerals within much younger rocks
3 109 ly3 (ly = light years). Yet it remains to from the oldest surviving terranes (Blichert-Toft
be seen whether the specific orbital architecture and Albarède 2008).
of our Solar System that includes habitable rocky That by 4.0 Ga the basaltic and the granitic
“▶ terrestrial planets” with liquid water within crust was established on Earth and oceans as well
a few astronomical units of a G-type star is excep- as a rock cycle and the generation of sediments is
tional or commonplace. directly supported by the existence of metamor-
Early Earth was a vastly different planet from phosed remnants of granitoid rocks and isotopic
the one we are so familiar with today. The sur- systems. Yet older evidence from the geochem-
face was heated from below by radioactivity, istry of detrital zircons (Jack Hills) tracks back to
thermal conduction through the ▶ crust, and vig- 4.4 Ga and shows that oceans and plate boundary
orous small-scale convection cells of ▶ magma processes (possibly plate tectonics?) could also
and mainly from above by the muted radiation of have been present in the Hadean (Hopkins
the young Sun and by occasional impacts. At et al. 2008). Hence, the old prevailing view of a
first glance, it should have been uninhabitable. largely molten surface blasted by sterilizing
However, if there were liquid water, organic impactors and intrinsically incapable of hosting
chemicals, and energy – all of which were life until after 3.8 Ga has been superseded by new
present on this primeval surface – it is not unrea- data. The new view of the Hadean eon
sonable to suppose that the primordial chilled (4.567–4 Ga; see ▶ Cool Early Earth) and subse-
crust was the first platform in which processes quent Eoarchean (4–3.6 Ga) favors by far an
leading to the emergence of life (▶ origin of life) earlier origin of life than had previously been
could have taken hold. Whatever the composi- thought (Fig. 1).
tion of the first crust was, the surface zone
remained under a super-greenhouse atmosphere Early Solar System, Earth, and Moon
that kept temperatures within the stability field The age of the Solar System is calculated to be
of liquid water. about 4.567 Ga (Connelly et al. 2008) based on
The early terrestrial hydrous surface was con- geochronological studies of the earliest solids
tinuously reworked by volcanic eruptions and that are native to the Solar System, which
Earth, Formation and Time 0

Sun and formation of
Early Evolution, Earth accretion, core formation since accretionary disk (4.57)
Fig. 1 Timeline of and degassing t0 (Myr)
Some differentiated
significant events in Earth’s Magma oceans on Earth asteroids (4.56)
100
early formation and Mars accreted (4.54)
Possible hot dense atmosphere
evolution Moon formation (~4.53)
Cooling of surface with loss of
dense atmosphere Loos of earth’s early
Opportunity life at this time? atmosphere (~4.53)
Earliest indications of felsic 200 Earth accretion, core

crust and liquid water on Earth formation, degassing,
complete (4.47)
Possibility of micro-continents
Earliest known zircon
fragment (4.38)
E
Environments conducive to
primitive life likely present 300 Upper age limit of most
hadean zircons (4.3)
Bombardments could have been
responsible for repeated destruction
of surface biosphere
400
Impact “bottleneck(s)” for the

early evolution of life?
Survival of life in a deep crustal

biosphere?
500
Earliest surviving
crust (4.02)
Stabilization of continental crust 600
and oceans
Late bombardment
of earth and moon (3.9)
Oldest known sediments,

700 Akilia, Greenland (~3.83)
Impact-induced selection for hyper-
Isua and Nuvvuagittuq
thermophilic life?
supracrustal belt
Earliest isotopic records implicating sediments (3.8–3.7)
biological activity
800 Warrawoona group (~3.5)
crystallized in the solar nebula before their even- solids. Direct astronomical observations of giant
tual incorporation into meteorites. The paradigm molecular clouds containing young stars show
for solar system formation begins with the col- that disk collapse and probable incipient dust
lapse of part of a giant molecular cloud (Safronov grain formation around such stars is common-
1969) that itself formed from gas and dust place (Pascucci et al. 2009). In the planet-
expelled in the explosion of one or more massive forming process, these grains aggregated into
stars. While the exact mechanisms responsible larger objects and eventually to planetesimals
for the initiation of nebular collapse are unclear, that were the building blocks of the planets.
nearby supernova explosions played a role in the A detailed chronology of the earliest events in
initial gravitational/mass instabilities of the cloud the Solar System, which span the time from the
based on the presence of the daughter products of first solids through the origin of the planets and
some extinct short-lived nuclides in the oldest ultimately to the formation of the ▶ crust,
▶ hydrosphere, and biosphere, will long be an hydrosphere recondensed out of the atmosphere
area of intense study. The physics of Earth accre- in less than 10 Myr (Sleep et al. 2001). It has been
tion has been explored in detail with sophisti- suggested that oceans were present on the proto-
cated dynamical computational tools that model Earth prior to the Moon-forming event (Abe
the interactions of planetesimals that led to the 1993). If there was a primary hydrosphere, it
growth (and in some cases the destruction) of was wholly vaporized by the ▶ giant impact and
ever-larger bodies. Numerical simulations of did not restabilize into a “secondary hydro-
terrestrial planetary growth succeed in modeling sphere” until later when the surface cooled to
the formation of planets of the size and distribu- the point where liquid water could begin to pool
tion as Mercury, Venus, Earth, and Mars on a thick (?) and dense primordial “chilled crust”
(Wetherill 1994). Such models show that most of likely ultramafic (Fe- and Mg-rich) composi-
of the mass of the Earth accreted in 10 Myr but tion at the top of the magma ocean (Arndt and
that there is an accretionary tail that extends for Chauvel 1991). It is possible that a primordial
an additional 100 Myr. Most of these dynamical crust and watery veneer of the post-impact
models seem to indicate that the colliding plan- Earth existed as early as about 35 million years
etesimals that built the terrestrial planets came after t-zero or around 4.53 Ga. A record of this
from beyond Mars’ orbit, or well beyond 2 AU primordial crust may never be found since all
(astronomical unit), and delivered volatile ele- traces of it are expected to have been destroyed
ments and compounds to the intrinsically dry, (cf. Carlson and Boyet 2009).
hot, inner Solar System well inboard from the Not until 150 Myr after the foundation of the
solar system’s “frost line” in the vicinity of the solar system (t0) do we have direct evidence of
asteroid belt. The climaxing event in the Earth’s the first (granitoid) crust from ancient terrestrial
formation and earliest evolution is the origin of zircons (at 4.38 Ga), and only 500 Myr after that
the Moon which was the result of the last big do we have the first direct evidence of surface
impact to affect the Earth. conditions from marine sediments and basaltic
The current paradigm favors the origin of the lavas (ca. 3.83 Ga). Hence, the geologic record
Moon no earlier than about 30 Ma into the history from actual rocks (as opposed to detrital zircons)
of the Solar System (i.e., from what is termed only extends back from the present through
“t-zero” at 4.567 Ga). Model studies indicate 88 % of Earth history. The Hadean zircons
that the Moon originated from a massive collision bring this record up to a remarkable 95 %. It is
of the proto-Earth when our planet was about not impossible that older actual rocks will be
90 % of its present mass (5.4 1024 kg) with found, but at present, the most ancient units are
another planet of about Mars’ mass slightly more than 4 Ga old and found in the
(6.5 1023 kg). Earth’s current mass is ▶ Acasta Gneiss Complex of northern Canada
5.974 1024 kg and that of the Moon is (Bowring and Williams 1999). A gallery of ages
7.35 1022 kg, so that it appears that mass loss for the next oldest rocks begins at the
from the impact process was negligible. The 3.77–3.87 Ga Isua Supracrustal Belt and the
impact was so energetic and deeply destructive ▶ Akilia association (Nutman et al. 1996) in
that it yielded a dense, hot, CO2-rock vapor- southern West Greenland and in Canada at the
steam atmosphere of hundreds of bars on top of ca. 3.75 Ga Nuvvuagittuq Supracrustal Belt in
a vigorously convective magma ocean that occu- northern Quebec (Cates and Mojzsis 2007). All
pied most of the Earth’s interior. An orbiting ring of these outcrops are dominated by highly
of super-hot devolatilized rock vapor coalesced deformed ▶ tonalite-trondhjemite-granodiorite
into the Moon (Canup and Asphaug 2001). The (TTG) gneisses that often (but not always) host
Moon-forming impact raised the whole Earth’s supracrustal enclaves of amphibolites and other
temperature to that of a small red star (>2,300 K) lithologies. Amphibolites in these cases are the
for several thousand years. For the neo-formed metamorphic equivalents of volcanic and intru-
Earth, most cooling models indicate that the sive igneous rocks such as basalt, gabbros, and
komatiites, and they sometimes host rare meta- these gases in the atmosphere. If all carbon cur-
morphosed sedimentary rocks such as ferrugi- rently present in Earth’s crust as carbonate and
nous siliceous schists (banded iron formations, organic matter was oxidized to CO2 and stored in
BIFs). Pillow lavas in the Isua belt unambigu- the atmosphere (Holland 1984), as happened on
ously attest to the presence of subaqueously Venus, the resulting 60–100 bar CO2 greenhouse
erupted basalts. could have produced surface temperatures to
180–230 C even as far back as 4.4 Ga (Abe
Early Atmosphere 1993). If pCO2 values were reduced to around
The Sun has increased its luminosity with time as 0.5 bar (still a 1,500-fold increase from modern
its core increased its density by hydrogen fusing values) by massive silicate weathering and car-
E
to helium. Standard models for the behavior of bonate sequestration in oceans as well as by
stars in the main sequence predict that the Sun enhanced volcanism and plate recycling, esti-
was only about 70 % as bright in the visible part mates for Hadean-Eoarchean surface tempera-
of the electromagnetic spectrum as it is now tures are reduced to near 0 C. Was the early
when it entered the main sequence 100 Myr Earth warm or cold? Controversy prevails over
after t0, around 4.46 Ga (Ribas et al. 2005). The the interpretation of proxy temperature data from
Sun had only brightened slightly more to 77 % isotopes of oxygen (Knauth 2005) and silicon
present value by the time of the earliest morpho- (Robert and Chaussidon 2006) in Precambrian
logical evidence of life in the form of oceans, temperature, and cherts. These data
bio-sedimentary structures (stromatolites) and have been interpreted to indicate seawater tem-
purported microbial microfossils which appear peratures above 50 C back to about 3.5 Ga.
much later in the rock record, about 3.5 Ga ago Atmospheric models constructed to simulate con-
in the Dresser Formation in Western Australia ditions on the early Earth are inconsistent with
(Van Kranendonk 2006). A dimmer early Sun the geochemistry of ancient sediments deposited
remains a long-standing and mostly unresolved in a hot ocean. Instead, model outputs for these
problem for atmospheric models that seek to atmosphere simulations almost always call for
keep the early Earth warm enough for liquid temperatures at or below the freezing point of
water to remain stable (Sagan and Mullen water. Hence, much more work remains to be
1972). So far, it is very difficult for these models done to resolve this fundamental discrepancy
to account for a warm early Earth, and no atmo- between model and experiment.
spheric models succeed in keeping ancient Mars
above 0 C. In the absence of abundant and Late Heavy Bombardment and Its Effects
efficient greenhouse gases, such as water vapor, During the epoch of heavy bombardment in the
CO2, CH4, and NH3, the Earth before about 2 bil- late Hadean and early Eoarchean (4.2–3.9 Ga),
lion years ago ought to have been ice covered to impacts modified geothermal, geochemical, and
equatorial latitudes. The only credible solution to geomorphologic conditions at the surface. Large
account for oceans of liquid water is to have a impactors, hundreds of kilometers in diameter,
denser atmosphere rich in powerful greenhouse can transform a globally habitable Earth into
gases compared to anything that has existed in a surface cauterized by a hundred meter thick
the Phanerozoic time; relatively speaking, there film of molten rock. However, only the largest
has been but a small variation in greenhouse of these impacts would have affected more than a
gases – well within several tens of percent of hemisphere, leaving the rest relatively
present atmospheric concentrations – over the unaffected.
last 500 Myr (Berner 1991). Almost all of our information about early
Broad estimates for average surface tempera- bombardment of the inner solar system comes
ture of the early Earth depend strongly on what from studies of the surface of the Moon. That
plausible limits can be placed on CO2, CH4, and object displays abundant evidence for an intense
NH3 abundances and the sources and lifetimes of impact flux at some time between the formation
of the lunar highland crust (ca. 4.5 Ga) and the of Earth’s mass). But uncertainty lingers over
outpourings of lava into the mare basins Earth’s total inventory of water (Mottl et al.
(>3.1 Ga). Ages of recovered fragments of the 2007), and the ultimate source of Earth’s water
ancient highland crust returned during the Apollo also remains ill constrained: it could be from
program have been interpreted to represent either intrinsic outgassing during planetary growth of
a short and intense “▶ late heavy bombardment” water supplied in accreting planetary material, or
(LHB) period at 3.9 Ga (Ryder 2002) or the tail it could have come subsequently from accreting
end of an extended post-accretionary bombard- meteoritic or cometary matter ▶ late veneer
ment (Hartmann et al. 2007). Some authors have (Albarède 2009). Probably all of these are cor-
argued that the absence of lunar impact melt ages rect. The availability and long-term stability of
older than about 3.9 Ga requires that the Solar liquid water over geologic timescales is probably
System was relatively quiet from the Moon- the determining factor that controls habitability
forming impact to the LHB (Ryder 2002). Inde- of a planet’s surface. Deciding whether Earth is
pendent verification of an LHB comes from stud- relatively “dry” or “wet” as terrestrial planets go
ies of lunar meteorites that record impacts from (Raymond et al. 2004) will depend on discoveries
the 3.9 Ga time frame (Cohen et al. 2000). If so, from future extrasolar systems by remote obser-
why was there an LHB? The exact mechanism is vations. Direct evidence for abundant surface
unknown, but early gravitational resonance inter- water on Earth appears to extend as far back as
action between Jupiter and the other outer planets the beginning of the geologic record.
has been implicated (Gomes et al. 2005). Mildly deformed and metamorphosed volca-
Although it is generally agreed that massive nic and sedimentary rocks that formed
collisions between Earth and space debris were subaqueously are found in 3.49 Ga rocks in the
fundamental formative agents to all of the Pilbara craton of Western Australia and the Bar-
planets in the first few hundred million years, berton greenstone belt of South Africa (Van
no clear terrestrial record of the LHB has so far Kranendonk 2006). However, all rocks older
been found (Anbar et al. 2001). Earth is a larger than about 3.6 Ga are highly deformed gneisses
body than the Moon, with stronger gravity to that have experienced pervasive metamorphism
attract bolides and more surface area to be and recrystallization. The most ancient gneissic
struck. By inference from the well-preserved terranes consist of more than 90 % of intrusive
lunar and Martian records, Earth’s crust during rocks and most of these are of granitoid (TTG)
the LHB ought to have been pummeled to dust. composition. These rock types, especially the
Although basic features of such an impact envi- tonalites and trondhjemites, are derived from
ronment have been investigated via numerical Na-rich and K-poor melts of hydrated mafic
models, the specific consequences of these col- (usually oceanic) crust following transformation
lisions for the crust and the overall effects they into garnet amphibolite or eclogite. As with most
had (destructive or procreative) to the emergent (but not all) present occurrences of TTGs, these
biosphere are an area of intense current study. rocks probably formed in subduction zones
Numerical modeling studies of the conse- (Martin et al. 2005). In older terranes such as the
quences to the Earth of impacting after the West ▶ Greenland Itsaq Gneiss Complex
Moon-forming event (ca. 4.53 Ga) show that (Nutman et al. 1996), multiple generations of
temperatures planetwide were never brought to granitoids of different ages and compositions
levels necessary for global sterilization host enclaves and tectonized rafts ranging in
(Abramov and Mojzsis 2009). size from meter to kilometer scale of yet older
“supracrustal” rocks which constitute something
Early Hydrosphere less than 10 % of the mapped outcrops. The oldest
Earth’s surface zone, from the top of the atmo- terrestrial rocks are found in the 4.03 Ga ▶ Acasta
sphere to mid-crustal depths, has more than one Gneiss Complex, but these deep-seated (7 km?)
ocean volume’s worth of liquid water (0.03 % intrusive igneous protoliths do not convey direct
information about the surface environment (Stern et al. 2007). Several workers have proposed a
and Bleeker 1998). Only when further discoveries minimum age of 3.83–3.85 Ga for the Akilia
are made of pre-3.6 Ga gneisses and supracrustal supracrustal rocks (Nutman et al. 1996; Man-
enclaves will more direct information be revealed ning et al. 2006), but this age, as well as the
about the early Earth. interpretation of these rocks as an originally
The oldest rocks with surviving volcanic and volcano-sedimentary succession, has been stri-
sedimentary structures indicative of a marine ori- dently challenged (Whitehouse et al. 2009).
gin form a tiny amount of the dominantly Other localities with >3.7 Ga volcanic or sed-
amphibolite (metamorphosed basalt) rocks of imentary rocks are found in the northeasternmost
the ca. 3.8 Ga Isua Supracrustal Belt which is margin of the Superior Province in the
E
part of the Itsaq Gneiss Complex. Within some ▶ Nuvvuagittuq Supracrustal Belt (previously
low-strain zones of the amphibolites, clear pillow known as Porpoise Cove). The Superior Province
structures are preserved, and because such struc- in northern Quebec is dominantly composed of
tures form when lavas erupt under water, they TTG and dioritic gneisses containing enclaves of
directly establish that liquid water was present older amphibolitized supracrustal rocks (Cates
when the Isua lavas erupted. At the Isua and Mojzsis 2009). As in West Greenland, these
Supracrustal Belt, associated with these rocks tend to comprise linear belts of metamorphosed
are quartz-magnetite schists that in places are sedimentary and extrusive igneous rocks that in
preserved as banded iron formations (BIF) with rare cases preserve primary sedimentary and vol-
genuine sedimentary banding; such sedimentary canic structures. Detailed mapping and geochro-
rocks also formed in liquid water (Klein 2005). nology of these units confirm a minimum age of
The Isua Supracrustal Belt contains the oldest 3.75 Ga for an 8 km2 enclave of mafic and ultra-
undisputed terrestrial record of a marine mafic amphibolites, quartz-magnetite BIFs,
environment. quartz-amphibole schists, and other possible
Of the other pre-3.8 Ga gneisses and amphib- detrital sediments at Nuvvuagittuq (Dauphas
olites in Greenland, none has been more closely et al. 2007).
scrutinized than the ca. 3.85 Ga-old outcrops on ▶ Plate tectonics’ mechanisms can be viewed
the island of Akilia (Manning et al. 2006). Rocks as an adequate, but not exclusive, means of
on ▶ Akilia include the polyphase ▶ Itsaq generating the ancient continental crust preserved
Gneiss Complex with ages that span in the West Greenland and northern Canada
3.87–3.62 Ga (Nutman et al. 1996). Unlike the terranes cited above. Alternatively, ▶ mantle
Isua rocks which encompass at least 200 km2 of superplumes (Albarède 1998) could have played
continuous exposure and preserve some primary at least a supporting role in the generation of this
structures, the gneisses on Akilia are very highly early crust. Future detailed studies of the oldest
deformed. Although the Akilia association is terranes will test the idea that plume’s plate tec-
widespread, with occurrences of enclaves tonics or some hybrid model can account for the
throughout the southernmost part of the Itsaq structure and geochemistry of the oldest pre-
Gneiss Complex, they are at much smaller served crust.
scale and are less coherent than the Isua belt.
At best, meter- to tens of meter-scale enclaves of Did Plate Tectonics Work Out in the Early
amphibolites and ferruginous quartzites are pre- Archean?
sent locally (Cates and Mojzsis 2006). Enclaves Within the most ancient terranes that have been
of Akilia quartz-magnetite-pyroxene schists mapped at the appropriate detail and which have
have been interpreted as metasediments, and been subjected to extensive geochronological
they share the trace element signatures of BIF studies, intrusive, volcanic, and sedimentary
from Isua and elsewhere (Manning et al. 2006). rocks of different ages are juxtaposed. Some
They preserve Fe isotope fractionations unlike authors have argued that this kind of geologic
igneous rocks and similar to Isua BIF (Dauphas style preserved in West Greenland is
a consequence of a compressive regime and plate • Conditions were conducive to the long-term
boundary processes that extended back to 3.8 Ga sustainability of biological activity.
or even earlier in the Hadean (Korenaga 2008; • Since the Hadean, actual preservation of
Davies 2008). Like modern oceanic arc evolu- Hadean zircons shows that Earth had crystal-
tion, many ancient but much better-preserved lized its magma ocean and formed stable crust
terranes, such as the 3.5 Ga Pilbara craton, record by 4.4 Ga and that no catastrophic impacts
permissive evidence of tectonic evolution from wholly destroyed that crust. Although the
plate- or plume-related crust formation to epi- nature of the primordial crust and the compo-
sodic growth of continental crust through the sition of the source rocks of the 4.4–4.0 Ga
emplacement of tonalites derived from partial zircons remain unresolved (Trail et al. 2007),
melting of hydrated basalt (Van Kranendonk there is now little doubt that the Hadean Earth
et al. 2007). The granitoids intruded sequences could have supported life.
of basalt with ferruginous-siliceous sedimentary
layers, a scenario remarkably similar in both Finally, although no rocks and minerals are
structure and chemistry to many contemporary preserved from Earth’s first 150 million years,
island arcs above subducting oceanic slabs there ought to be echoes of its prior existence.
(Kato et al. 1998). Recent work from both Nd and Hf isotope signa-
tures from 4.4 to 3.5 Ga rocks and minerals pro-
Did Plate Tectonics Exist in the Hadean? vides evidence of planetary-scale events during
Zircons older than 3.9 Ga are mostly found at the Earth’s first 100 million years. These events
Jack Hills and Mt. Narryer in the Narryer Gneiss that entailed major chemical fractionations in the
Complex, Yilgarn craton, Western Australia mantle, perhaps as a consequence of an early
(Froude et al. 1983; Compston and Pidgeon magma ocean and the establishment of the
1986). Others have been recovered from else- proto-crust in the early Hadean, occurred prior
where in Western Australia (Wyche et al. 2004) to about 4.3 billion years ago or even earlier
and in scattered localities worldwide (e.g., (Caro et al. 2006; Blichert-Toft and
Mueller et al. 1998). Many of the most ancient Albarède 2008; Tessalina et al. 2010).
Jack Hills’ zircons contain inclusions of quartz,
muscovite, and other phases produced from
silica-saturated igneous rocks, meaning that See Also
granites existed before 4.1 Ga (Hopkins
et al. 2008). Low temperatures of formation, con- ▶ Acasta Gneiss
sistent with plate underthrusting, have been ▶ Akilia
determined using titanium thermometry on ▶ Archean Environmental Conditions
>4.0 Ga Jack Hills’ zircons. Many Hadean zir- ▶ Archean Tectonics
cons also have isotopically heavy oxygen (d18O ▶ Chondrite
up to +7.3 ‰) which supports the view that melt- ▶ Condensation Sequence
ing of hydrated ▶ crust produced the pre-4.0 Ga ▶ Continental Crust
granites and that some were possibly contami- ▶ Cool Early Earth
nated by subducted sediment (Mojzsis ▶ Craton
et al. 2001). Although the details of this hypoth- ▶ Earth’s Atmosphere, History of the Origins
esis remain the focus of debate, the viewpoint of a ▶ Earth’s Atmosphere, Origin and Evolution of
hospitable, water-rich ▶ Hadean Earth leads to a ▶ Earth, Age of
number of predictions: ▶ Exoplanets, Discovery
▶ Geochronology
• Separation of crust into continental (granitic) ▶ Geological Time Scale, History of
and oceanic (mafic) styles prevailed since the ▶ Giant Impact
earliest times. ▶ Greenstone Belts
▶ Hadean rocks: constraints on the early differentiation of the

▶ Isua Supracrustal Belt Earth’s mantle. Geochim Cosmochim Acta 70:164–191
Cates NL, Mojzsis SJ (2006) Chemical and isotopic evi-
▶ Jack Hills (Yilgarn Craton, Western Australia) dence for widespread Eoarchean metasedimentary
▶ Komatiite enclaves in southern West Greenland. Geochim
▶ Late Heavy Bombardment Cosmochim Acta 70:4229–4257
▶ Late Veneer Cates NL, Mojzsis SJ (2007) Pre-3750 Ma supracrustal
rocks from the Nuvvuagittuq supracrustal belt, north-
▶ Late-Stage Accretion ern Quebec. Earth Planet Sci Lett 255:9–21
▶ Mantle Plume, Planetary Cates NL, Mojzsis SJ (2009) Metamorphic zircon, trace
▶ Mantle Volatiles elements and Neoarchean metamorphism in the
▶ Nuvvuagittuq (Porpoise Cove) Greenstone ca. 3.75 Ga Nuvvuagittuq supracrustal belt, Quebec
(Canada). Chem Geol 261:98–113 E
Belt Cohen BA, Swindle TD, Kring DA (2000) Support for the
▶ Ocean, Chemical Evolution of lunar cataclysm hypothesis from lunar meteorite
▶ Oceans, Origin of impact melt ages. Science 290:1754–1756
▶ Oxygen Isotopes Compston W, Pidgeon RT (1986) Jack Hills, evidence of
more very old detrital zircons in Western Australia.
▶ Plate Tectonics Nature 321:766–769
▶ Prebiotic Chemistry Connelly JN, Amelin Y, Krot AN, Bizzarro M (2008)
▶ Radiogenic Isotopes Chronology of the solar system’s oldest solids.
▶ System Solar Formation, Chronology of Astrophys J Lett 675:L121–L124
Dauphas N, Cates NL, Mojzsis SJ, Busigny V (2007)
Identification of chemical sedimentary protoliths
using iron isotopes in the >3750 Ma Nuvvuagittuq
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Nutman AP, McGregor VR, Friend CRL, Bennett VC, Champion DC (2007) Review: secular tectonic evolu-
Kinny PD (1996) The Itsaq gneiss complex of southern tion of Archean continental crust: interplay
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early crustal evolution (3900–3600 Ma). Precambrian mation of the Pilbara Craton, Australia. Terra Nova
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Neodymium-142 evidence or hadean mafic crust. Sci- Geochim Cosmochim Acta 58:4513–4520
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Pascucci I, Apai D, Luhman K, Henning T, Bouwman J, Controversy: field, structural and geochronological
Meyer MR, Lahuis F, Natta A (2009) The different evidence questions interpretations of >3.8 Ga life in
evolution of gas and dust in disks around Sun-like and SW Greenland. J Geol Soc 166:335–348
cool stars. Astrophys J 696:143–159 Wyche S, Nelson DR, Riganti A (2004) 4350–3130 Ma
Raymond SN, Quinn T, Lunine JI (2004) Making other detrital zircons in the Southern Cross Granite-
Earths: dynamical simulations of terrestrial planet for- Greenstone Terrane, Western Australia: implications
mation and water delivery. Icarus 168:1–17 for the early evolution of the Yilgarn Craton. Aust
Ribas I, Guinan EF, Gudel M, Audard M (2005) Evolution J Earth Sci 51:31–45
of the solar activity over time and effects on planetary
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strom). Astrophys J 622:680–694
isotopes in cherts. Nature 443:969–972 Earth, Surface Evolution
Ryder G (2002) Bombardment of the Hadean earth:
wholesome or deleterious? Astrobiology 3:3–6 Nicholas Arndt
Safronov VS (1969) Evolution of protoplanetary cloud
and the formation of the Earth and Planets. Moscow,
Nauka (in Russian); NASA Tech. Transl. F-677,
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http://exoplanet.eu/ rocks; South Africa; Traces of life
Earth, Surface Evolution 699
Definition Archean cherts suggest that ocean temperatures

were commonly above 40 C (Knauth and Lowe
The surface of the Archean Earth was in many 2003). The atmosphere contained a little to no
ways similar to that of today. Oceans covered free oxygen but was rich in CO2; rainwater
most of the globe, but there were also regions of was acid.
dry land. Oceanic crust was almost as thick as The normal cycle of erosion, transport, and
continental crust, mountain ranges were not deposition of sediment operated, but the rivers
very high, and parts of oceanic ridges and pla- flowed through a landscape that was very differ-
teaux were emergent. Geological processes ent from that of today. The feature that most
such as volcanism, erosion, and sediment starkly distinguished the Archean and modern
E
deposition operated as now but were influenced land surface was the lack of vegetation.
by a lack of vegetation, higher ocean tempera- Microbes no doubt colonized the subsurface
tures, and a hotter more aggressive, acidic and constructed biofilms (slime mats) that cov-
atmosphere. ered moist areas, but most of the landscape
was a Martian vista of bare rocks and soil. The
rate of erosion was enhanced by the lack of
Overview vegetation, high temperatures, and aggressive
atmosphere but restrained by modest heights of
The total area covered by oceans was greater mountain belts. Active volcanism covered much
than now, for two reasons. First, the volume of of the surface with lava flows or pyroclastic
continental crust may have been less, if this crust deposits.
had grown progressively through time (Benn The ▶ oceanic crust was composed of basaltic
et al. 2006). Second, the oceans were more volu- lavas like that of modern crust but more magne-
minous because high temperatures in the mantle sian (picritic) in places (Sleep and Windley
(Nisbet et al. 1993) destabilized hydrous min- 1982). Parts of mid-ocean ridges and the summits
erals and drove water to the surface. Mountain of oceanic plateaux may have been emergent
ranges existed but were not as high as those of forming what might be called “melano (dark-
today because the continental crust was heated colored)-continents.” The pelagic sediment that
internally and rendered more ductile by more covered this crust was different from that of
abundant radioactive elements. Continental today. An absence of shell-forming organisms
crust was relatively thin, while oceanic crust, precluded the formation of biogenic calcareous
produced by high-degree melting of the hotter or siliceous oozes; in their place were Si- or
mantle, was far thicker (Sleep and Windley Fe-rich sediments that precipitated directly from
1982). The subdued topography, the limited con- the high-temperature seawater that contained
trast between the thicknesses of oceanic and high concentrations of these elements. Hydro-
continental crust, combined with bigger oceans, thermal circulation of Si-charged seawater
meant that much of the continental crust was resulted in massive silicification of all near-
flooded (Arndt 1998). surface rocks. Expulsion of fluids at hydrother-
Just as during more recent geological history, mal vents led to the deposition of exhalative
global temperatures waxed and waned. Periods sediments variably composed of sulfides, sul-
of global glaciation, the most pronounced being fates, or carbonates or silica minerals (Russell
during the Proterozoic “▶ snowball Earth” epi- et al. 2005).
sodes (Hoffman et al. 1998), alternated with The earliest Archean coincided with the end of
periods when temperatures were relatively the ▶ Late Heavy Bombardment, a time of mas-
high. The O and Si isotopic compositions of sive meteorite impacts. The largest of these
700 Earth’s Atmosphere, History of the Origins
would have vaporized large expanses of the Sleep NH, Zahnle K (2001) Carbon dioxide cycling and
oceans and pulverized large parts of the conti- implications for climate on ancient Earth. J Geophys
Res 106:1373–1399
nents, but their overall impact was local, not Valley JM, Peck WH, King EM, Wilde SA (2002) A cool
global. early Earth. Geology 30:351–354
Opinions differ concerning the surface envi-
ronment in the Hadean. One school envisages
a ▶ cool early Earth in which clement condi-
tions reigned (Valley et al. 2002); another
imagines a hellish environment in which Earth’s Atmosphere, History of the
periods of intense heating subsequent to mete- Origins
orite impact alternated with global glaciation
when the atmosphere budget of greenhouse Stéphane Le Gars
gases was insufficient to counteract the weak Centre François Viète, Université de Nantes,
luminescence of the young sun (Sleep and Nantes, France
Zahnle 2001).
Keywords
See Also
Ampère; Atmosphere; Kant-Laplace cosmo-
▶ Archean Traces of Life gonic hypothesis; Koene; Lavoisier; Rubey;
▶ Earth, Formation and Early Evolution Vernadsky
▶ Impact Melt Rock
▶ Komatiite History
▶ Pilbara Craton If the scientific investigation of the atmosphere
▶ Plate Tectonics really started in the seventeenth century, the
problem of its origin was taken into account
only two centuries later. It is indeed with the
works of Galileo Galilei, Blaise Pascal, and
Arndt NT (1998) Why was flood volcanism on submerged Robert Boyle that the notion of pressure and
continental platforms so common in the precambrian? temperature permitted to study in a quantitative
Precambrian Res 97:155–164 way the atmosphere. John Mayow, Boyle’s stu-
Benn K, Mareschal J-C, Condie KC (2006) Archean dent, concluded, in an empirical way, that the air
geodynamics and environments. Geophysical mono-
graph series. Am Geophys Union 164:320 is a two-body mixture out of which one takes
Hoffman PF, Kaufman AJ, Halverson GP, Schrag DP place at once in breathing and burning. In 1777,
(1998) A neoproterozoic snowball earth. Science Antoine-Laurent de Lavoisier initiated an analy-
281:1342–1346 sis and synthesis of air, and found out the “aerial
Knauth LP, Lowe DR (2003) High Archean climatic tem-
perature inferred from oxygen isotope geochemistry of nitrogen” and “oxygen” terms to name the two
cherts in the 3.5 Ga Swaziland supergroup, South gases of the atmosphere. In his memoir “Vues gé
Africa. Geol Soc Am Bull 115:566–580 nérales sur la formation et la constitution de
Nisbet EG, Cheadle MJ, Arndt NT, Bickle MJ l’atmosphère,” Lavoisier even thought of the pos-
(1993) Constraining the potential temperature of the
Archaean mantle: a review of the evidence from sibility of the existence, at a high altitude, of
komatiites. Lithos 30:291–307 a hydrogen film, which would sometimes be
Russell MJ, Hall AJ, Boyce AJ, Fallick AE (2005) On destroyed completely by combustion and would
hydrothermal convection systems and the emergence form again by the continual decomposition
of life. Econ Geol 100:419–438
Sleep NH, Windley BF (1982) Archaean plate tectonics: of plant and animal matters generated at the
constraints and inferences. J Geol 90:363–379 earth’s surface.
Earth’s Atmosphere, Origin and Evolution of 701
In 1833, the French physicist André-Marie References and Further Reading

Ampère drew the conclusions from Laplace’s
hypothesis concerning the solar system origin Allègre C, Staudacher T, Sarda P (1986/1987) Rare gas
systematics: formation of the atmosphere, evolution
and, taking up the Lavoisier’s point of view and
and structure of the Earth’s mantle. Earth Planet Sci
expressing the idea that the earth’s atmosphere Lett 81:127–150
results from a reduction of bodies oxygenated by Aristote (2008) Météorologiques. Flammarion, Paris,
metals, leading to a nitrogen release originally pp 138–139
Arrhenius S (1929) Le destin des étoiles. Alcan, Paris, p 118
combined to oxygen, explaining also the origin of
Dalmas J-B (1852) La cosmogonie et la géologie, basées
oxygen in the atmosphere. As far back as 1856, the sur les faits physiques, astronomiques et géologiques
Belgian chemist Corneille Jean Koene considered qui ont été constatés ou admis par les savants du
dix-neuvième siècle, et leur comparaison avec la for-
E
a link between oxygen and carbonic acid gas,
mation des cieux et de la terre selon la genèse.
inferring the lack of oxygen gas in the original
Imprimerie de Louis Perrin, Lyon, p 188
terrestrial atmosphere because of a reducing Dauphas N (2003) The dual origin of the terrestrial atmo-
medium due to the occurrence of metal or sulphide sphere. Icarus 165:326–339
compounds. Finally, in the 1920s, the Russian Dauvillier A (1947) Genèse, Nature et Évolution des
planètes. Cosmogonie du système solaire. Géogénie.
Wladimir Vernadsky introduced, by his biosphere
Genèse de la vie. Hermann, Paris, p 154
concept, the idea of a balance between atmosphere Doel RE (1996) Solar system astronomy in America:
and human beings: for him, gases of the atmo- communities, patronage, and interdisciplinary science,
sphere are in an equilibrium state of dynamic and 1920–1960. Cambridge University Press, Cambridge
Flammarion C (1873) L’atmosphère. Description des
perpetual exchange with living matter.
grands phénomènes de la nature. Hachette, Paris,
It is only in 1951 that American geologist Wil- pp 38–39
liam Rubey in a seminal paper (Geologic history Laplace PS, Exposition du système du monde, Livre IV,
of sea water: An attempt to state the problem) that chapitre 9
Lavoisier Antoine Laurent de (1777) Analyse de l’air de
a modern theory of the origin of the atmosphere
l’atmosphère, sa résolution en deux fluides élastiques,
shaped up. Rubey proposed that the Earth’s atmo- l’un respirable, l’autre non respirable. Antoine-
sphere did not originate from the weathering of Laurent de Lavoisier
the continental crust, a view that was popular Lavoisier Antoine Laurent de, Vues générales sur la for-
mation et la constitution de l’atmosphère
among geologists at that time, but rather origi-
Rubey W (1951) Geologic history of sea water: an
nated from volcanic degassing throughout geolog- attempt to state the problem. Geol Soc Am Bull 62:
ical times. The development of the noble gas 1111–1147
geochemistry in that period supported this view.
Von Weizäcker (1937) clearly understood that the
gaseous argon-40 was a product of degassing of
the solid potassium-40 which resides in the crustal Earth’s Atmosphere, Origin and
rocks. Analyses of radiogenic nuclides of neon, Evolution of
argon, and xenon in the 1980s (Allègre et al. 1986/
1987) allowed quantifying a chronology of the Bernard Marty
degassing and demonstrated that majority of Institut Universitaire de France, Ecole Nationale
atmospheric gases did not derived from primor- Supérieure de Géologie, Centre de Recherches
dial gases from a primitive solar nebula but rather Pétrographiques et Géochimiques (CRPG),
added shortly later the formation of the Earth by CNRS, Vandoeuvre les Nancy Cedex, France
meteoritic and/or cometary material (Dauphas
2003).
Keywords
See Also
Atmosphere; Carbon; Comets; Mantle volatiles;
▶ Earth’s Atmosphere, Origin and Evolution of Meteorites; Nitrogen; Noble gases; Water
702 Earth’s Atmosphere, Origin and Evolution of
Definition dissociated to release CO2, the resulting atmo-

spheric pressure would be around 27 bars, and
The Earth’s atmosphere is a thin gaseous layer the CO2/N2 ratio would be comparable to those of
made of oxygen, ▶ nitrogen, and trace gases. Venus and Mars atmospheres. Oceanic water,
It permits a mild temperature and provides a which is liquid under the Earth’s surface condi-
protection against cosmic rays, solar wind, and tions but was probably gaseous during the forma-
meteorite falls. It was stable enough through tion of the Earth, was part of the early
geological time to have nurtured the develop- atmosphere. If oceanic water was vaporized and
ment of life. Originally, the atmosphere oceanic halogens (components of salt)
consisted of water, carbon dioxide, nitrogen, transformed into their gaseous forms (HCl, HF,
sulfur, and ▶ noble gases, and it evolved by HBr), then the atmospheric pressure would be
condensation of water to form the oceans, trap- 220 bar instead of 1.01 bar, and the Earth’s sur-
ping of ▶ carbon in carbonates and organic mat- face would be an inferno hostile to life. The
ter, and biogenic production of oxygen. The reason why Earth escaped such hellish fate
Earth’s atmosphere originated probably from (while Venus kept CO2-rich atmosphere with
the addition of bodies from the asteroid belt a pressure of 90 bar and a surface temperature
between Mars and Jupiter toward the end of of 450 C) is unclear and is the focus of active
terrestrial formation. research.
Overview Origin of the Terrestrial Atmosphere
The Earth’s atmosphere consists mainly of oxy- The “reconstructed” atmosphere’s composition,
gen (20.95 % O2) and nitrogen (78.078 % N2), the that is, the sum of volatile elements in the air, in
other constituents making only 1 % of the total the oceans, and in sedimentary rocks, consists of
volume: argon (0.934 %), CO2 (preindustrial: 86 % water vapor, 12 % CO2, 1.4 % HCl + HF,
280 ppm vol/vol), and noble gases. This compo- and 0.2 % N2. This is quite comparable to the
sition is unique among planetary atmospheres: composition of volcanic gases emitted by active
atmospheres of the giant planets (Jupiter, Saturn, volcanoes. This observation led Williams
Uranus, Neptune) are dominated by hydrogen W. Rubey, a geologist at the US Geological
and helium, reflecting the composition of the Survey, to propose that the Earth’s atmosphere
protosolar nebula, whereas the atmospheres of did not originate from weathering (chemical
the inner planets, Venus and Mars (Mercury alteration) of the continental crust, a view that
does not possess an atmosphere), are rich in was popular among geologists at that time, but
CO2, the concentration of N2 being of the order rather originated from volcano degassing
of a few %. The high oxygen content of the throughout geological times (Rubey 1951). In
terrestrial atmosphere is due to life. Oxygen is support of this view was the discovery of the
produced by photosynthesis, a mechanism that decay of 40K (a rare isotope of potassium) pro-
uses solar photons to break down and oxidize ducing 40Ar, the most abundant isotope of the
organic molecules. Without life, oxygen noble gas argon. In fact, the 0.934 % abundance
would disappear in a few thousand years due to of Ar in the air is almost entirely due to 40Ar, the
iron oxidation in terrestrial rocks. The very low other more abundant argon isotope, 36Ar, having
CO2 content of air is the result of trapping of an atmospheric concentration of only 32 ppm
carbon in rocks, either as carbonates or as organic vol. The discoverer of the 40K decay, Von
matter buried in the crust. If carbonates were Weizsäcker, a German physicist in pre-World
War II Germany, noted correctly that the high that could have fractionated noble gas isotopes
40
Ar content of air was the result of the (Pepin 1991). In particular, the isotopic compo-
▶ degassing of the “solid,” silicate-rich Earth, sition of mantle neon is close to that observed in
where potassium resides (Von Weizäcker 1937). the solar nebula (Sarda et al. 1988), which sug-
This key observation led to the concept of atmo- gests that part of volatile elements originated
spheric formation by degassing of volatile ele- from the nebular gas or from matter implanted
ments trapped in the mantle. This concept was by ions from the young Sun. This problem is not
refined using noble gases. Some of the isotopes yet resolved, but some insight arises from the
of the chemically inert elements, now in the stable isotopes (i.e., the isotopes that are not
atmosphere, were produced by natural nuclear produced or destroyed by nuclear reactions) of
E
reactions like radioactive decay and spontane- water and nitrogen. The ratios of the two hydro-
ous fission of parent elements contained in the gen isotopes from the water molecule, D/H
mantle and crust, thus establishing a chronology (where D stands for deuterium, or 2H, and
of atmospheric formation. The terrestrial mantle H for 1H), and of the two isotopes of atmospheric
also contains volatile elements including noble nitrogen, 15N/14N, are comparable to values
gases. A comparison of noble gas isotopic com- found in primitive meteorites, also called chon-
positions between the mantle (which can be sam- drites (Marty et al. 2010). They are markedly
pled in basalts originating from mantle melting) different from the isotopic signatures of the
and the atmosphere shows that the latter formed cloud of gas and dust that collapsed to form the
very early in the Earth’s history, within Solar System and from isotopic ratios measured
100 Ma after the start of solar system forma- in comets (another potential source of atmo-
tion, 4.57 Ga ago (Ozima 1975; Staudacher and spheric volatiles, since comets, which come
Allègre 1982). from the outer solar system beyond the orbit of
There is evidence that a source other than Neptune, are very rich in volatile elements with
mantle degassing contributed to atmospheric water making up to 50 %). Hence, major vola-
formation. On the one hand, models of ▶ solar tiles like water and nitrogen, and probably car-
system dynamics suggest that the Earth formed bon too, seem to originate mostly from bodies
in a region of the nascent Solar System too hot to within the inner Solar System, akin of present-
permit trapping of volatile elements into plane- day asteroids between Mars and Jupiter. This
tary embryos. Therefore, it has been proposed scenario is consistent with dynamical models
that terrestrial volatiles might have been contrib- advocated above in which volatile-rich asteroi-
uted directly by impacting of bodies originating dal bodies contributed to Earth toward the end of
away from the present-day Earth radial location, accretion. These bodies were not necessarily
in a region between Mars and Jupiter where big, and gas-rich dust like interplanetary parti-
volatile-rich meteorites, called the carbonaceous cles or micrometeorites could have played a sig-
chondrites, originate. From dynamical reasons, nificant role (Maurette et al. 2000). Mass
these contributions might have happened toward balance considerations involving siderophile as
the end of the terrestrial accretion once more well as lithophile elements indicate that the con-
than 90 % of the Earth was formed (Morbidelli tribution in volatile-rich material at the end of
et al. 2000). On the other hand, the noble gas accretion could have made only 0.3 % of the
isotopic composition of ▶ mantle volatiles is total terrestrial mass.
significantly different from that of the atmo- Atmospheric noble gas isotopes indicate also
sphere, which suggests either different cosmo- that the early atmosphere had been escaping to
chemical sources for the mantle and the space for prolonged periods of time of the order
atmosphere (Marty 1989) or drastic atmospheric of several hundreds of million years, as a result of
processing such as escape of volatiles to space intense UV irradiation by the early Sun, which
704 Earth’s Atmosphere, Origin and Evolution of
dissociated the water molecules (Hunten living matter, if any), but such effect is presently
et al. 1987). Meanwhile, large bodies battered unknown.
the Earth, resulting in large-scale melting and The composition of the early atmosphere is
vaporization. Such high temperature episodes constrained by the rock record. The composition
did not last for more than 1 or 2 Myr due to of ancient lavas called ▶ komatiites indicates
radiative heat loss into space, and, at some that their oxygen fugacity (a measure of the
stage, the water vapor started to condensate to oxidation state of a mineral assemblage) was
form the oceans. According to the isotopic record close to that of modern lavas (Canil 2002). The
of some rare, very ancient, ▶ zircons, liquid gas phase in equilibrium with such a molten
water might have existed as soon as 4.37 Ga magma contains water vapor, CO2, and N2 as
ago, that is, only 150 Ma after the start of the main species of H, C, and N. Hence, the
terrestrial accretion. Besides water, the Earth’s early atmosphere of the Earth in the Hadean
surface needed to get rid of CO2 also: this species, (4.4–3.8 Ga) was probably oxidizing and not
the likely prevalent form of carbon in the early reducing as proposed from experiments aimed
atmosphere, is a strong greenhouse gas that, if to produce amino acids in a reduced gas mixture
abundant, could have prevented cooling of the upon electric discharges (Miller 1953). How
Earth’s surface to the condensation temperature oxidizing the atmosphere was during the first
of water. How this took place is unclear: carbon 1–2 Ga is not clear yet. Sulfur in sedimentary
might have been efficiently returned to the mantle rocks of the Archean eon (the geological eon
by subduction (Zahnle 2006), or it could have between 3.8 Ga ago and 2.5 Ga ago) contains
formed carbonates, providing that liquid water isotopic anomalies that are best explained by
was available and Ca2+ ions were removed by interactions between atmospheric species and
surface alteration of basalts. ultraviolet irradiation of the Sun. At present
It is probable that, before 3.8 Ga ago, large and presumably since 2.4 Ga, most of solar UV
ocean volumes already existed, as indicated by irradiation is absorbed in the higher atmosphere
the occurrence of sedimentary rocks in geologi- by oxygen species, notably ozone. Hence, the
cal formations of this age. The carbon isotopic evidence from sulfur isotopes that UV photons
composition of graphitic inclusions, encapsu- could penetrate deep in the atmosphere has been
lated in apatite in quartz from 3.86 Ga-old taken as evidence that free oxygen was not pre-
metasedimentary rocks from ▶ Akilia, West sent in the atmosphere until 2.4 Ga ago
Greenland, suggests that life already existed (Farquhar et al. 2000).
(Mojzsis et al. 1996), but this record is still During the first half of the Earth’s history, the
controversial. Around 3.8 Ga ago, a ▶ Late Sun was less luminous, and the energy reaching
Heavy Bombardment (LHB) of extraterrestrial the terrestrial surface was 25 % smaller than
material is recorded at the lunar surface by Maria the present-day one (Kasting 1993). However,
craters dated at 3.85–3.85 Ga (Tera et al. 1974). no glaciation took place in the Archean eon (the
Scaled to the Earth’s dimensions, the LHB so-called ▶ faint young Sun paradox), and there
would have contributed an average thickness of is evidence that the oceans were hotter, up to
about 200 m of extraterrestrial matter onto the 70 C on average, than the present-day oceans
whole Earth’s surface. Modeling of solar system (Robert and Chaussidon 2006; ▶ Precambrian
dynamics suggests that the LHB was due to Oceans, Temperature of). More CO2 in the
a large-scale perturbation of the outer Solar Sys- Archean atmosphere might have helped in keep-
tem and that both comets and asteroidal bodies ing the Earth’s surface warm, but the required
were involved (Gomes et al. 2005). This episode level is beyond what the sedimentary record
had certainly significant impact on the atmo- from this time has preserved in terms of CO2
spheric composition (and on the survival of partial pressure (e.g., Shaw 2008). The existence
of methane in the Archean atmosphere requires ▶ Archean drilling projects are sampling
a strong source, and the only significant one unaltered rocks for this purpose.
could be CH4-producing bacteria. Alternatively,
it has been proposed that more N2 in the atmo-
sphere would have helped to preserve a signifi- See Also
cant greenhouse effect in the Archean, by
increasing Rayleigh backscattering (Goldblatt ▶ Carbon
et al. 2009). There is no consensus about the ▶ Faint Young Sun Paradox
time of appearance of free oxygen in the atmo- ▶ Great Oxygenation Event
sphere, a strong indication of colonization of the ▶ Mantle Volatiles
E
Earth’s surface by living matter. Different lines ▶ Nitrogen
of evidence point to a (geologically) brutal ▶ Nitrogen Isotopes
increase of the partial pressure of O2 around ▶ Noble Gases
2.35 Ga ago, the Great Oxygenation Event ▶ Ocean, Chemical Evolution of
when life-related sources of oxygen took over ▶ Oceans, Origin of
oxygen consumption by basalts (Holland 2002), ▶ Oxygenation of the Earth’s Atmosphere
and the isotopic signature of sulfur in sedimen- ▶ Stardust Mission
tary rocks discussed above seems to support
this view.
Canil D (2002) Vanadium in peridotites, mantle redox and

Future Directions tectonic environments: archean to present. Earth
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atmosphere will arise in the near future. First, son AJ, Zahnle KJ (2009) Nitrogen-enhanced green-
several planetary missions are documenting house warming on early Earth. Nat Geosci 2:891–896
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Origin of the cataclysmic late heavy bombardment
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Meteorit Planet Sci 35:1309–1320 significant earth-like atmospheres and which
Ozima M (1975) Ar isotopes and Earth-atmosphere evolu- orbit, e.g., different classes of main sequence
tion models. Geochim Cosmochim Acta 39:1127–1140 stars or/and which are at different positions in
Pepin RO (1991) On the origin and early evolution of the habitable zone.
terrestrial planetary atmospheres and meteoritic vola-
tiles. Icarus 92:1–79
isotopes in cherts. Nature 443:969–972
Rubey WW (1951) Geologic history of sea water. Geol
Soc Am Bull 62:1111–1148 Earth-Like Planet
Sarda P, Staudacher T, Allègre CJ (1988) Neon isotopes in
submarine basalts. Earth Planet Sci Lett 91:73–88 ▶ Terrestrial Planet
Shaw GH (2008) Earth’s atmosphere – Hadean to early
Proterozoic. Chem Erde-Geochem 68:235–264
Staudacher T, Allègre CJ (1982) Terrestrial xenololy.
Earth Planet Sci Lett 60:389–405
Tera F, Papanastassiou D, Wasserburg G (1974) Isotopic
evidence for a terminal lunar cataclysm. Earth Planet Eccentricity
Sci Lett 22:1–21
Von Weizäcker CF (1937) Über die Möglichkeit eines
dualen-Zerfalls von Kalium. Phys Z 38:623–624 Daniel Rouan
Zahnle KJ (2006) Earth’s earliest atmosphere. Elements LESIA, Observatoire Paris-Site de Meudon,
2:217–222 Meudon, France
Earth’s Oxygenation Definition
▶ Oxygenation of the Earth’s Atmosphere The eccentricity is a parameter (denoted by e)

characterizing the orbit’s shape for an object
gravitationally bound to another one. It quan-
tifies the amount by which the orbit deviates
Earth-like Atmosphere from a circle: e = 0 for a circle, e = 1
for a parabola, e > 1 for a hyperbola, and
John Lee Grenfell 0 < e < 1 for an ellipse, where e = c/a,
German Aerospace Center (DLR), Berlin, with c the distance from the center to a focus
Germany and a the semimajor axis. See Fig. 1 for
an illustration of different values in the case
of an ellipse. The eccentricities of planets
Definition belonging to the solar system are rather small,
while a significant fraction of the discovered
An earth-like atmosphere is one whose dom- exoplanets exhibit large eccentricities, in
inant constituents are molecular nitrogen and excess of 0.3.
Eclipse 707
Eccentricity,
Fig. 1 Several examples
of elliptical orbits with e=0
different eccentricities (circle)
e = 0.23
e = 0.43
e = 0.77
passes in front of the hotter one. This is tradition-

Eclipse ally denoted by the primary eclipse. The second-
ary eclipse refers to the shallower event when
David W. Latham1 and Nader Haghighipour2 the cooler component passes behind the
1
Harvard-Smithsonian Center for Astrophysics, hotter one. However, there can be interesting
Cambridge, MA, USA exceptions to this general pattern. For example,
2
Institute for Astronomy, University of Hawaii- with highly eccentric orbits and modest inclina-
Manoa, Honolulu, Hawaii, HI, USA tions, one or the other of the eclipses may not
occur.
By convention, the eclipse of a star by a much
Synonyms smaller and cooler object such as a planet is
called a ▶ transit, while the passage of the planet
Eclipsing binary; Primary eclipse; Secondary behind the star is termed either an occultation or a
eclipse; Transit secondary eclipse. Another interesting case is a
binary consisting of a normal star and a ▶ white
dwarf. White dwarfs are small enough to produce
Definition transit-like events when they pass in front of the
star. But, if they are young enough, they can be
In an astronomical system, when orbits of two or hotter than the star, thus producing a deeper
more objects are viewed edge-on, the compo- eclipse when they pass behind the star.
nents periodically pass in front of each other, Eclipsing binaries for which the spectra of
thus reducing the amount of light that reaches both stars can be detected are especially valuable.
the observer. This phenomenon is known as In these systems, an analysis of the light curve
Eclipse. If both components are stars, the system (the graphs of the variation of the intensity of the
is called an eclipsing binary. Normally the light received from the system in term of time)
deeper eclipse occurs when the cooler object together with the spectroscopic orbit derived
708 Eclipsing Binary
from the radial velocities of the two stars can the middle of the zodiacal belt that extends about
yield the absolute mass and radius of each stellar 9 on either side and which contains the orbits of
component. If the spectrum of only one of the all the major planets orbiting the Sun. By exten-
stars is detected (presumably that of the brighter sion, the ecliptic is often assimilated to the average
one), then only the ratios of the masses and radii plane of the solar system. The term ecliptic also
can be derived directly. Interestingly, the density designates the adjective that specifies quantities
of the primary star and surface gravity of an associated with the ecliptic, for example, ecliptic
unseen companion can also be derived from the coordinates.
light curve and orbital solution. If the mass of the
primary can be estimated from other observa-
tions, such as spectroscopy and/or parallaxes See Also
combined with stellar models, then the actual
mass and radius of the secondary can be derived. ▶ Coordinate Systems
Transiting planets are simply an extreme exam- ▶ Orbit
ple of the case where only the spectrum of the ▶ System Solar Formation, Chronology of
primary star yields an orbital solution. ▶ Zodiacal Light
See Also
▶ Activity, Magnetic Ecological Niche

▶ Transit
▶ Transiting Planets Felipe Gomez
▶ White Dwarf Centro de Astrobiologı́a (CSIC/INTA), Instituto
Nacional de Técnica Aeroespacial, Torrejón de
Ardoz, Madrid, Spain
Eclipsing Binary Definition

▶ Eclipse
Niche in ecology is a term which refers to the
relational position of a species or population in an
ecosystem (environment). It refers to the behav-
ior of the population to any external input or
Ecliptic
relationship including resources, competitors,
physical conditions, etc.
Daniel Rouan
Meudon, France
Ecology, History of
Definition Stéphane Tirard

The ecliptic is the geometric plane containing the des Techniques EA 1161, Faculté des Sciences et
Earth’s ▶ orbit. More precisely, it is the average des Techniques de Nantes, Nantes, France
plane of the orbit of the center of mass of the Earth-
Moon system. It is also the line delineating the
projection of the Earth’s orbital plane onto The word ecology comes from the ancient Greek
the celestial sphere. On the sky, the ecliptic is at oikos, house, and logos, science. It was coined by
Ecosystem 709
E. Haeckel in 1866 and means the relationship interdependent biological (biotic) components,
between living beings and their environment. in which the community of living organisms con-
Toward the end of the nineteenth century, this tinually interacts with all abiotic components
relation became a research topic for botanists. For (e.g., rock, water, air, soil, sunlight) through var-
instance, in 1895, Eugen Warming was a pioneer ious interconnected relationships. The flow of
of the study of the distribution of plants. energy within an ecosystem, manifested through
In 1935, G. Tansley created the word ecosys- cycling of materials between the biotic and abi-
tem, which he defined as the functional ecologi- otic components, defines its inherent trophic
cal unit. During the 1950s, E.P. and H.T. Odum, structure and biological diversity.
inspired by thermodynamics, developed central
E
concepts, such as a theory of open systems and
the notion of a dynamical equilibrium regulated Overview
by energy flows. That was the development of
systemic ecology. The ecosystem concept developed, as a more
abstract replacement of the concept of commu-
nity, out of theorizing in the 1930s on the orga-
nization and dynamics of natural systems.
Ecopoesis While a community is a group of populations
of different organisms that interact with one
▶ Planetary Ecosynthesis another in a given ▶ habitat or area and are
interdependent, an ecosystem is considered an
area where living communities exchange mate-
rials “. . .with the whole complex of physical
factors that form what we term an environment”
Ecosphere (Tansley 1935). Ecosystems are therefore
essentially self-contained energy and nutrient
▶ Biosphere cycles. An ecosystem can be long lived or tem-
porary, and balance is a critical element of any
viable ecosystem because disturbances and
instability in its physical factors (e.g., rapid
Ecosystem climate change) can threaten the existence of
its component organisms. Ecosystems can be
J. Cynan Ellis-Evans small or large, and the physical boundaries of
UK Arctic Office, Strategic Coordination Group, an ecosystem can be defined as in a pond, or
British Antarctic Survey, Cambridge, UK relatively blurred, as in the case of a marsh
draining into a stream or river. Ecosystems are
everywhere and the diversity is enormous on
Keywords a planet such as Earth, and indeed we are
still discovering environments (e.g., the deep
Abiotic; Biogeochemical cycles; Biological subsurface, subglacial lakes) that harbor as
populations; Biotic; Environment; Physicochem- yet barely researched ecosystems. Extreme
ical conditions environments on Earth provide analogues of
possible extraterrestrial ecosystems. Potential
locations include subsurface permafrost litho-
Definition sols on Mars; the ice-covered ocean of
the Jovian moon, Europa; the hydrocarbon
An ecosystem is part of the broad ▶ environment, lakes on Titan; or the subsurface water on
comprising of both physical (▶ abiotic) and Enceladus.
710 ECSS
See Also
Ediacara Biota
▶ Abiotic
▶ Biogeochemical Cycles ▶ Ediacaran Biota
▶ Environment
▶ Habitat Ediacaran Biota
Thomas H. P. Harvey
Department of Earth Sciences, University of
Costanza R, D’Arge R, de Groot R, Farber S, Grasso M, Cambridge, Cambridge, UK
Hannon B et al (1997) The value of the world’s eco-
system services and natural capital. Nature
387:253–260. doi:10.1038/387253a0 Keywords
Tansley AG (1935) The use and abuse of vegetational
terms and concepts. Ecology 16:284–307
United Nations Environment Programme. Convention on Evolution; Paleobiology; Precambrian
biological diversity. June 1992. UNEP document No
Na.92-78
Synonyms
Ediacara biota
ECSS
Catharine A. Conley Definition

A particular suite of fossil organisms of the Edi-
acaran Period of geologic time.
Definition
The European Cooperation for Space Standard- Overview

ization (ECSS) is, since 1993, an organization
that works to improve standardization within the This somewhat loose term (sometimes Ediacara
European space sector. This organization is or Ediacaran biota) is widely used to refer to the
supported by the ▶ European Space Agency macroscopic, often complex, soft-bodied and
and the space agencies of some member states. notably problematic marine organisms that are
Industry is also participating in the technical characteristic of the later part of the Ediacaran
working groups which define the standards to Period (ca. 575–542 Ma), although there is poten-
be applied to the management, the organization, tial confusion with a broader usage referring to
the quality assurance of the projects, as Ediacaran-age life in general. The biota is named
well as the engineering methods in various after the Ediacara Hills region, South Australia,
domains (electrical system, communication, but almost 40 localities are now known world-
mechanic, etc.). wide, with further key assemblages described
from England, Russia, Namibia, and Newfound-
land, Canada. Evidence for the origins and fate of
See Also the biota is sparse, although a rather earlier
(>635 Ma) assemblage of simple forms from
▶ European Space Agency northwestern Canada has been reported, as have
▶ ISO some possible Cambrian “survivors.”
ee 711
The Ediacaran biota includes the earliest References and Further Reading
known fossils of conspicuously large and com-
plex organisms. Because this occurrence is pre- Knoll AH (2003) Life on a young planet: the first three
billion years of evolution on Earth. Princeton Univer-
ceded by a long period of apparently slow-paced
sity Press, Princeton
microbial evolution and succeeded by the sudden Narbonne GM (2005) The Ediacara biota: neoproterozoic
appearance of diverse and unambiguous meta- origin of animals and their ecosystems. Annu Rev
zoan (animal) fossils of the Cambrian explosion, Earth Planet Sci 33:421–442
Vickers-Rich P, Komarower P (eds) (2007) The rise and
the phylogenetic identities and ecological capa-
fall of the Ediacaran biota. Geological Society special
bilities of the Ediacaran organisms are of pivotal publication 286. The Geological Society, London
significance. At first, the tendency was to inter- Xiao S, Laflamme M (2009) On the eve of animal radia-
tion: phylogeny, ecology and evolution of the Ediacara
E
pret them as members of various familiar meta-
biota. Trends Ecol Evol 24:31–40
zoan groups, such as cnidarians, echinoderms,
and annelids. Subsequently, however, their often
bizarre-looking morphologies prompted the pro-
posal of alternative phylogenetic positions rang-
ing across the tree of life. Those of modular Ediacaran Period
construction were suggested to constitute a dis-
tinct (and extinct) group termed the Vendobionta. Thomas H. P. Harvey
Increasingly, the Ediacaran biota is being viewed Department of Earth Sciences, University of
as a collection of probably rather diverse organ- Cambridge, Cambridge, UK
isms, perhaps including some true members of
extant metazoan lineages (such as the putative
bilaterian Kimberella) alongside members of
Definition
extinct stem lineages to more inclusive groups
(including possible stem metazoans), with others
The period of geologic time between c. 635 and
potentially identifiable as protists, algae, fungi,
542 Ma and the youngest period of the Protero-
and microbial colonies.
zoic Eon. The upper boundary marks the base
The ecologies of the Ediacaran organisms are
of the Cambrian Period and the beginning of
often challenging to interpret. Some, notably
the Phanerozoic Eon. The lower boundary is
Kimberella, appear to be preserved alongside
defined by the upper limit of Marinoan glacia-
feeding traces. Most, however, have no clear
tion rocks in Enorama Creek, Australia. Fossils
capabilities for movement or ingestion, and
from the Ediacaran Period provide the earliest
some lived at too great a depth in the oceans to
abundant evidence for complex, large-bodied
have been photosynthetic. The forms with mod-
organisms, notably in the distinctive Ediacaran
ular construction show adaptations consistent
biota.
with osmotrophy, the direct absorption of nutri-
ents across body surfaces. Interpreting the phy-
logeny, ecology, and evolutionary dynamics of
the Ediacaran biota remains a key challenge in See Also
paleobiology.
▶ Ediacaran Biota
▶ Proterozoic Eon
See Also
▶ Ediacaran Period
▶ Eukaryotes, Appearance and Early Evolution ee
of
▶ Proterozoic Eon ▶ Enantiomeric Excess
712 Effective Temperature
temperature of 225 K, in order to maintain the

Effective Temperature balance between the absorbed solar flux and the
emitted thermal radiation. Note that the effective
Daniel Rouan temperature depends on the reflectivity, or
LESIA, Observatoire Paris-Site de Meudon, albedo, of the planet: the larger the ▶ albedo,
Meudon, France the lower the effective temperature.
Definition See Also
The effective temperature of a star is the temper- ▶ Blackbody

ature of a ▶ blackbody of the same radius as the ▶ Bolometric Magnitude
star and that would radiate the same total amount ▶ Color Index
of electromagnetic power as the star. ▶ Emissivity
▶ Grey Body
▶ Hertzsprung-Russell Diagram
Overview
Noted Teff, the effective temperature is one of

the fundamental parameters that characterizes a EH
star. If P* is the power radiated by a star of
radius R, then Teff is derived by applying the ▶ Redox Potential
Stefan-Boltzmann law: P ¼ 4 p R2 s Teff 4 .
Indeed a star is not actually a blackbody, as its
▶ emissivity varies with wavelength; however,
the effective temperature generally provides a Ejecta
fair approximation of the actual temperature of
the stellar photosphere. This is so because at any Daniele L. Pinti
wavelength, the radiation comes from a layer GEOTOP Research Center for Geochemistry and
which is always on the skin of the star (the Geodynamics, Université du Québec à Montréal,
▶ photosphere) and, thus, at approximately a Montréal, QC, Canada
constant temperature. The effective temperature
is directly related to the color of the star: the
higher the temperature, the bluer the light emit- Synonyms
ted by the star. The effective temperature is one
of the two parameters, with the stellar luminos- Pyroclastics (in volcanology)
ity, used to build the classical HR diagram that
permits astronomers to classify stars. The effec-
tive temperature of the Sun is 5,780 K, while it is Definition
40,000 K for an O star and 3,000 K for an M star.
The term has been extended to planets, with a Ejecta is solid, liquid, or gaseous material ejected
similar definition, but, there, the effective tem- from a source region. Three different kinds of
perature can sometimes be fairly different from ejecta exist: impact ejecta, volcanic ejecta
the surface temperature. This is, for instance, the (or pyroclastics), and (stellar) ejecta. Impact
case of Venus, where the ▶ greenhouse effect in ejecta is solid, liquid, or vaporized rock debris
the atmosphere (due to CO2) blocks the infrared ejected ballistically from a ▶ crater during a
outward radiation imposing a ground tempera- meteoritic or cometary impact. These ejecta can
ture of 735 K, much higher than the effective be dispersed around the crater and form specific
Electric Discharge 713
patterns or partially build the crater rim. The See Also

volcanic ejecta is rock debris ejected by an explo-
sive ▶ volcano and normally grouped in the more ▶ Escape Velocity
general term pyroclastics. In astrophysics, ejecta ▶ Exoplanets, Discovery
is the material expelled from a ▶ star, either ▶ Orbit
quiescently (e.g., in coronal mass ejection or ▶ Planet Formation
through the wind of an evolved star, like a ▶ red ▶ Planetesimals
giant or an AGB star) or in a stellar explosion like
in a ▶ supernova or nova.
References and Further Reading E
Rasio FA, Ford EB (1996) Dynamical instabilities and the
See Also formation of extrasolar planetary systems. Science
274:954–956
▶ Breccia Weidenschilling S, Marzari F (1996) Gravitational scat-
tering as a possible origin for giant planets at small
▶ Crater, Impact stellar distances. Nature 384:619–621
▶ Impactite
▶ Red Giant
▶ Star
▶ Supernova Electric Discharge
▶ Volcano
Jeffrey Bada
CA, USA
Ejection, Hyperbolic
Synonyms
Sean N. Raymond
Spark discharge
Definition
Definition
An electric discharge is the release and transmis-
Hyperbolic ejection is the removal of an object sion of electricity in an applied electric field
from a gravitational system (e.g., a planetary through a medium such as a gas. Several types
system) on a hyperbolic ▶ orbit, so that the of electric discharges occur naturally on Earth
object is permanently lost from the system. (American Geophysical Union 1986):
This is caused by an increase in a body’s orbital
energy that exceeds the gravitational binding 1. Atmospheric lightning, which is thought to be
energy of the system. In planetary systems, the caused by the frictional generation, and sepa-
increase in orbital energy usually comes from a ration, of positive and negative charges on ice
close encounter with a massive planet. Objects and dust particles. As the charge on these
that are ejected from planetary systems are particles builds up, the result is the often-
thought to include a large number of ▶ planetes- spectacular discharge of electricity known as
imal-sized bodies (asteroids and comets), a lightning. On average, there are 50–100 light-
small number of ▶ planetary embryo-sized bod- ning strikes per second on Earth with most of
ies, and, in some cases, fully grown terrestrial, the activity taking place in equatorial and
ice giant, or gas giant planets. northern latitudinal regions.
714 Electrochemical Potential
2. Corona discharges, which are caused by an Lightning in the solar nebula may have also
electrical discharge produced by the ionization been important in the synthesis of reduced spe-
of the surrounding atmosphere, generating a cies of importance in prebiotic chemistry.
luminous plasma (sometime referred to as Electric discharges were first used in the con-
St. Elmo’s fire, a term used by sailors to text of astrobiology research in the 1952 Miller-
describe the glow observed at the top of a Urey Experiment (Miller 1953) in which a spark
ship’s mast during a thunderstorm). Unlike was passed through a mixture of methane, ammo-
lightning, which is instantaneous and transient, nia, hydrogen, and water, simulating the action of
coronal discharges are less brilliant, can last for lightning acting on what was then thought to be
a significant period of time (minutes or more), the composition of the primitive atmosphere.
and occur over a large surface area. Continuous This experiment ultimately yielded amino acids,
corona discharges have been observed on the suggesting a facile route from simple environ-
nose cones of airplanes during thunderstorms. mental processes to the building blocks of mod-
3. Volcanic lightning, which is ubiquitous in most ern organisms.
volcanic eruptions, especially water-rich erup-
tions. The cause of the lightning is not See Also
completely understood but is thought to be the
charging of ash particles by frictional processes ▶ Ozone
and the ionization of gases expelled in the erup- ▶ Spark Discharge
tion. Volcanic lightning has been observed to
be nearly continuous during an eruption.
American Geophysical Union, National Research Council

Overview (U.S.). Geophysics Study Committee (1986) The
earth’s electrical environment. National Academy
Electrical discharges have also been observed in Press, Washington, DC, p 263
the atmospheres of other bodies in the solar system Aplin KL (2006) Atmospheric electrification in the solar
system. Surv Geophys 27:63–108
(Aplin 2006). Lightning has been detected during Cleaves HJ, Chalmers JH, Lazcano A, Miller SL, Bada JL
observations of Jupiter and Saturn, and the NASA/ (2008) A reassessment of prebiotic organic synthesis
ESA Cassini/Huygens mission has tentatively in neutral planetary atmospheres. Orig Life
detected lightning in the atmosphere of Titan. Evol Biosph 38:105–115
Miller SL (1953) A production of amino acids under possible
Electric discharges on Earth, and by implica- primitive earth conditions. Science 117(3046):528–529
tion elsewhere, play an important role in various
aspects of atmospheric chemistry. On the
present-day Earth, the major products of light-
ning acting on the gases in the atmosphere are Electrochemical Potential
NO, along with lesser amounts of ▶ ozone. On
the primitive Earth, a possible reducing atmo- Juli Peretó
sphere containing nitrogen and methane would Institut Cavanilles de Biodiversitat i Biologia
have had HCN as the major product. In contrast, Evolutiva, Universitat de València, València, Spain
with a neutral nitrogen-carbon dioxide-rich pri-
mordial atmosphere, the major product would
have been a mixture of NO and CO, as well as Definition
ammonia and HCN (Cleaves et al. 2008).
In the hydrogen-rich Jovian atmosphere, the Electrochemical potential is the free energy change
products produced by lightning are much more (DG) associated with the movement of chemical
diverse and include, besides HCN, low molecular species through a gradient of electric potential
weight hydrocarbons and formaldehyde. and/or a gradient of concentration established
Electron Acceptor 715
between the two sides of a biomembrane. The

electrochemical potentials of H+ and/or Na+ are Electromagnetic Spectrum
a main fuel for the cell energy transductions.
Daniel Rouan
See Also
Meudon, France
▶ ATP Synthase
▶ ATPase
Definition
▶ Bioenergetics
▶ Photosynthesis E
The electromagnetic spectrum is the domain of all
possible frequencies or wavelengths over which
▶ Respiration
electromagnetic waves can be observed. This is a
▶ Transduction
huge domain spanning 23 decades in frequencies or
wavelengths. It includes radio waves, millimetric
waves, infrared, visible light, ultraviolet, x-rays,
Electromagnetic Radiation and gamma rays. The distribution versus frequency
(or wavelength) of the radiation intensity emitted
Daniel Rouan by a particular celestial object is called its electro-
LESIA, Observatoire Paris-Site de Meudon, magnetic spectrum or spectral energy distribution
Meudon, France (SED); it contains in general extremely rich pieces
of information on the nature and physical proper-
ties of the object, such as temperature, composition,
Definition velocity, etc. ▶ Spectroscopy is the art of measur-
ing the spectrum of a source.
The expression electromagnetic radiation desig-
nates the energy that is transported in the form of See Also
a propagating wave made of two perpendicular
components, one an electric field and the other a ▶ Electromagnetic Radiation
magnetic field, which oscillate at the same fre- ▶ Spectroscopy
quency. This description applies to light in all its
forms: radio waves, infrared, visible light, ultravi-
olet, x-rays, and gamma rays. It is by far the most
important vector of information reaching us that Electron Acceptor
comes from the universe and the objects it con-
tains. Different physical processes give rise to Juli Peretó
electromagnetic radiation and are generally classi- Institut Cavanilles de Biodiversitat i Biologia
fied as thermal (resulting from the thermal motion Evolutiva, Universitat de València, València, Spain
in a medium), nonthermal radiation (synchrotron),
or quantized (transitions between levels of energy
Synonyms
in atoms, molecules, or nuclei).
Oxidant; Oxidative agent
See Also
▶ Blackbody Definition
▶ Bremsstrahlung Radiation
▶ Radiative Processes Electron acceptor is any of the organic or inor-
▶ Synchrotron Radiation ganic oxidative agents participating in an
716 Electron Attachment
enzymatic reaction. In a restrictive sense, it is the See Also

terminal oxidant of an electron transport chain.
▶ Coenzyme
See Also ▶ Cofactor
▶ Cytochromes
▶ Anaerobic Respiration ▶ Electron Acceptor
▶ Photosynthesis ▶ Electron Donor
▶ Respiration ▶ NADH, NADPH
▶ Photosynthesis
▶ Respiration
Electron Attachment
Steven B. Charnley
Electron Dissociative Recombination
Steven B. Charnley
Space Flight Center, Greenbelt,
MD, USA
Definition
Electron attachment is the chemical process

Definition
whereby an electron is attached to a neutral mol-
ecule. Examples include electron photo attach-
Electron dissociative recombination is the chemi-
ment to CN to produce CN and a photon and
cal process whereby an electron reacts with a pos-
dissociative electron attachment to HCN to pro-
itive molecular ion to produce two neutral species.
duce CN and a hydrogen atom.
Examples include electron recombination with
H3O+ to produce OH and 2H. This is a very impor-
tant process in gas-phase interstellar chemistry.
See Also
▶ Interstellar Chemical Processes See Also
Electron Carrier
Juli Peretó
Electron Donor
Juli Peretó
Definition
Spain
An electron carrier is any of the soluble (e.g.,
NADH) or protein-bound (e.g., ▶ cytochromes) Synonyms
▶ cofactors participating as reversible ▶ electron
donor and/or acceptor in an enzymatic reaction. Reducing agent; Reductant
Electron Transport 717
Definition
Electron Transport
Electron donor is any of the organic or inorganic
substances acting as reducing agents in an enzy- Francisco Montero
matic reaction. In a restrictive sense, the reducing Department of Biochemistry and Molecular
agent at the beginning of an ▶ electron transport Biology I, Facultad de Ciencias Quı́micas,
chain or the substrate is used to obtain energy by Universidad Complutense de Madrid, Madrid,
▶ respiration. Spain
E
See Also Keywords
▶ Electron Transport Electron carriers; Photosynthesis; Redox reac-

▶ NADH, NADPH tions; Respiration
▶ Photosynthesis
▶ Respiration
Synonyms
Electron transport chain

Electron Radiative Recombination
Definition
Steven B. Charnley
Electron transport refers to the transfer of elec-
trons from an initial donor (reducing substance)
to a final acceptor (oxidizing substance) across
different intermediaries, which takes place in
biological membranes. Electron transport pro-
Synonyms
cesses are usually associated with respiratory
and photosynthetic processes.
Radiative recombination
Overview
Definition
The most common ways by which living organ-
Electron radiative recombination is a chemical
isms obtain energy from their surroundings is by
process whereby an electron reacts with a posi-
the ▶ oxidation of external electron donors
tive atomic ion to produce a neutral atom with the
(chemotrophic organisms) or by the absorption
emission of a photon (i.e., radiation). Examples
of electromagnetic radiation (phototrophic
include electron reaction with H+ to produce a
organisms). In both cases, the energy translation
neutral H atom plus a photon; photons produced
mechanisms take place in membranes (the cyto-
in this process are responsible for the red color of
plasmic membranes in the case of prokaryotes or
many low-density nebulae.
subcellular particles such as mitochondria and
chloroplasts in the case of eukaryotes). In the
See Also translation process, an electron transport takes
place from a substance, which is an initial
▶ Electron Dissociative Recombination ▶ electron donor to a final acceptor, and since
▶ Interstellar Chemical Processes the ▶ redox potential of the donor is more
718 Electron Transport Chain
negative than that of the acceptor, the global ▶ Photosynthesis

process is exergonic. This oxidation-reduction ▶ Redox Potential
process takes place across a series of intermedi- ▶ Reduction
aries (▶ Electron Carrier), which are not free in ▶ Respiration
the intracellular medium as the majority of them
form highly structured complexes inserted in the
membranes. Therefore, the transfer of electrons
that takes place between the different compo- Nelson DL, Cox MM (2005) Lehninger principles of bio-
nents does not occur using the conventional chemistry, 4th edn. W.H. Freeman, New York
mechanism that takes place in the oxidation- Nicholls DG, Ferguson SJ (2002) Bioenergetics, 3rd edn.
reduction reactions of a homogenous system in Academic, London
Voet D, Voet JG (2004) Biochemistry, 3rd edn. Wiley,
dissolution. Instead, a directional electron trans- New York
port takes place along the membrane and, in White D (1999) The physiology and biochemistry of pro-
some cases, a transverse one as well. This karyotes, 2nd edn. Oxford University Press, New York
explains how energy coupling between an
oxidation-reduction reaction and a transport
phenomenon is possible, as first, the free energy
released by these processes is used to produce
a proton transport against the gradient across the Electron Transport Chain
membrane, and then the proton ▶ electrochemi-
cal potential gradient is translated, by means of ▶ Electron Transport
processes that also take place in the membrane,
in ▶ ATP.
For many years, the mechanism whereby elec-
trons are transferred between the different elec-
tron carriers, which are frequently separated from Electrophoresis
each other when the structure of the complexes is
greater than 10 Ȧ, has been the subject of much Adrienne Kish
debate. However, there is growing experimental Institut de Génétique et Microbiologie,
and theoretical evidence that suggests that on Universite Paris-Sud 11, Orsay Cedex, France
some occasions the transport may be brought
about by the “tunnel effect.”
The molecules that take part in the electron Definition
transport are quite ubiquitous. The most common
molecules are cytochromes, quinones, iron-sulfur Electrophoresis is the migration of charged mol-
proteins, and flavoproteins. In addition, chloro- ecules (for example ▶ nucleic acids or ▶ pro-
phylls and pheophytins are involved in teins) through a medium according to charge
▶ photosynthesis. and molecular size as a consequence of an
applied electric current and interactions with
the medium. It is used as an analytical technique
See Also to separate molecules by charge, size, or binding
affinity. Parameters including matrix composi-
▶ ATP tion and concentration, ionic strength, electrical
▶ Electrochemical Potential current strength and field angle, and migration
▶ Electron Acceptor time can be varied. Electrophoretic methods
▶ Electron Carrier commonly employed include capillary electro-
▶ Electron Donor phoresis (CE), gel electrophoresis (GE), and
▶ Oxidation pulsed-field gel electrophoresis (PFGE) for
Eley-Rideal Mechanism 719
separation of chromosomal DNA; denaturing History

and/or temperature gradient gel electrophoresis
(DGGE/TGGE) for separation according to The definition of “cosmic abundances” also takes
sequence; polyacrylamide gel electrophoresis account of the relative abundances in carbona-
(PAGE) and SDS-PAGE for separation of pro- ceous chondrites, which generally correlate well
teins in their native and denatured conforma- with those in the solar photosphere (Anders and
tions, respectively; and isoelectric focusing Grevesse 1989; Przybilla et al. 2008) apart from
(IEF) and 2D electrophoresis for separation of the most volatile elements such as hydrogen and
proteins by isoelectric point with or without helium.
PAGE first.
E
See Also
See Also
▶ Abundances of Elements
▶ DNA Sequencing ▶ Carbonaceous Chondrite
▶ Nucleic Acids ▶ Diffuse Cloud
▶ Protein ▶ Interstellar Dust
Elemental Carbon Anders E, Grevesse N (1989) Abundances of the

elements – meteoritic and solar. Geochim Cosmochim
Acta 53:197–214
▶ Carbon Przybilla N, Nieva M-F, Butler K (2008) A cosmic abun-
dance standard: chemical homogeneity of the solar
neighborhood and the ISM dust-phase composition.
Elemental Depletion
Steven B. Charnley
Solar System Exploration Division, Code 691, Elephant Trunks
Space Flight Center, Greenbelt, MD, USA ▶ Pillars
Definition
Eley-Rideal Mechanism
Elemental depletion denotes the difference in
elemental composition of an astronomical William M. Irvine
source relative to some standard of reference, University of Massachusetts, Amherst, MA, USA
usually the Sun. Lines of sight through the
local interstellar medium indicate much lower
abundances of many chemical elements com- Definition
pared to the solar composition. This is largely
due to the differential incorporation of these The Eley-Rideal mechanism is a theoretical
elements into interstellar dust grains. Different model of how some bimolecular chemical reac-
lines of sight through diffuse clouds show dif- tions can take place on solid surfaces. In the
ferent depletions. theory, one atom or molecule is adsorbed on the
720 Embden-Meyerhof-Parnas Pathway
surface, and another reacts directly from the gas History

phase. This may be contrasted with the
▶ Langmuir-Hinshelwood mechanism. In the By 1940, the canonical glycolytic pathway (i.e.,
interstellar medium, the formation of molecular the one responsible for alcoholic fermentation in
hydrogen is thought to occur primarily through yeasts and anaerobic ▶ glycolysis in muscle) was
reactions on the surfaces of dust grains. elucidated. Actually, it was the result of a collec-
tive task with the contributions of, among others,
Gustav Embden (1874–1933), Arthur Harden
See Also (1865–1940), Karl Lohmann (1898–1978), Otto
Fritz Meyerhof (1841–1951), Jakub Karol Parnas
▶ Adsorption (1884–1949), and Otto Heinrich Warburg
▶ Interstellar Dust (1883–1970).
▶ Langmuir-Hinshelwood Mechanism
See Also
▶ Entner-Doudoroff Pathway
Embden-Meyerhof-Parnas Pathway ▶ Fermentation
▶ Gluconeogenesis
Juli Peretó
▶ Metabolism
Spain
Embedded Bioburden
Synonyms
▶ Encapsulated Bioburden
Glycolysis
Definition
Emergence of Life
The Embden-Meyerhof-Parnas (EMP) pathway
▶ Origin of Life
allows the metabolic use of glucose to generate
ATP, NADH, and several biosynthetic precur-
sors such as 3-phosphoglycerate or pyruvate.
The EMP pathway can occur both anaerobi-
cally (leading to one or several ▶ fermentation Emission Nebula
pathways) and aerobically through the conver-
sion of pyruvate to acetyl CoA and the connec- William M. Irvine
tion with the tricarboxylic acids (TCA) cycle. University of Massachusetts, Amherst, MA, USA
The classical version of the EMP pathway is
present in bacteria and eukaryotes whereas sev-
eral modified versions are present in anaerobic Definition
archaea. The second half of the pathway is
almost universal, and thus, it could represent That portion of an interstellar cloud that is heated
the oldest part of the pathway, related to a and ionized by radiation from nearby stars shines
primordial origin of ▶ gluconeogenesis. as an emission nebula. The spectrum is
Enantiomeric Excess 721
characterized by emission lines from the constit-

uent atoms and ions. Emission nebulae are typ- 100 % Enantiomeric Excess
ically H II regions, although planetary nebulae
are sometimes also referred to as emission ▶ Homochirality
nebulae.
See Also
Enantiomeric Excess
▶ HII Region
E
▶ Planetary Nebula Robert Hazen
Geophysical Laboratory, Carnegie Institution of
Washington, Washington, DC, USA
Emissivity
Synonyms
Daniel Rouan
Chiral excess; ee
Meudon, France
Definition
Definition
Enantiomeric excess (ee) is a measure of the
deviation of a mixture of ▶ chiral (i.e., handed)
Emissivity is a quantity in physics that depends
molecules from the equimolar (or “racemic”)
on wavelength and that characterizes the effi-
state. Enantiomeric excess is usually expressed
ciency with which a surface radiates, compared
as a percent; if D and L are the amounts of two
to a ▶ black body at the same temperature.
chiral molecules, the enantiomeric excess is
A black surface has an emissivity close to 1 and
given as
a polished reflecting surface an emissivity close
to 0. Because of Kirchhoff’s law, the same quan-
ee ¼ ½ðD LÞ=ðD þ LÞ 100
tity characterizes also the ability of a surface to
absorb or to reflect impinging radiation
Thus, the ee of a 60:40 mixture of D:L is 20 –
(absorptivity = emissivity, ▶ albedo = 1 emis-
a value that can be thought of as a mixture with
sivity). Emissivity is a positive, dimensionless
20 % pure D plus 80 % racemic mixture; further-
quantity, lower than or equal to 1.
more, ee equals 0 and 100 for racemic and
chirally pure mixtures, respectively.
An enantiomeric excess can be measured in a
See Also
number of ways, including specific optical rota-
tion or chromatography with a chiral column. The
▶ Albedo
enantiomeric excess is important in characteriz-
▶ Blackbody
ing the purity of chiral pharmaceuticals and other
chiral products. Many models of the origins of
life postulate the local and/or global development
of enantiomeric excesses in mixtures of amino
Empire acids or other biomolecules and their subsequent
chiral amplification, as a prelude to life’s
▶ Domain (Taxonomy) ▶ homochirality.
722 50:50 Enantiomeric Mixture
An enantiomeric excess has been reported

in ▶ carbonaceous chondrites for some organic
compounds, including amino acids (see
▶ Chirality).
See Also
▶ Amino Acid
▶ Chirality
▶ Enantiomers
▶ Homochirality
50:50 Enantiomeric Mixture
▶ Racemic Mixture
Enantiomers, Fig. 1 An amino acid (bottom) showing

four different atoms (R is another carbon substituent, e.g.,
CH3) attached to a central carbon atom. This configura-
Enantiomeric Ratio tion results in the property of asymmetry. The resulting
enantiomers are mirror images of each other [much like
▶ D/L-Ratio the mirror images of your hands (top)], but they are oth-
erwise chemical equivalent
have identical physical and chemical properties,

Enantiomers with the exception that they rotate light in equal
but opposite directions. Enantiomers based on
Jeffrey Bada other central atoms such as silicon, phosphorus,
Scripps Institution of Oceanography, La Jolla, and germanium are also known.
CA, USA
See Also
Definition
▶ Amino Acid
▶ Chirality
An enantiomer is one of a pair of optical ▶ iso-
▶ Stereoisomers
mers, the structures of which are not superimpos-
able on their mirror images. In organic
compounds, enantiomers contain a carbon atom
that has four different groups attached to it
(asymmetric or chiral carbon) (see Fig. 1). The Enantiopure
best-known examples are the L- and
D-enantiomers of amino acids. Enantiomers ▶ Homochirality
Enceladus 723
Definition
Encapsulated Bioburden
Enceladus is one of the midsized icy satellites of
Catharine A. Conley ▶ Saturn. It was discovered by William Herschel
NASA Headquarters, Washington, DC, USA in 1789. Its distance to Saturn is 238,100 km or
about four Saturnian radii. Its diameter is 500 km
and its density is 1.0 g/cm3, corresponding to
Synonyms water ice. With an albedo of 0.9, Enceladus is
the brightest object known in the solar system;
Embedded bioburden such a high albedo results in very low surface
E
temperature (70 K).
Definition
Overview
In planetary protection, the term “encapsulated
▶ bioburden” is used to indicated the number of The exploration of Enceladus started with the Voy-
viable microorganisms that are trapped inside non- ager missions. The first images of Voyager 1,
metallic spacecraft materials. They can be trapped taken in December 1980, already indicated a bright
between the layers of multilayer insulator, in the and young surface with relatively few craters.
glue for honeycombs, or in the matrix itself. Voyager 2, in August 1981, not only confirmed
A spaceflight project may choose to measure the this result but also provided evidence for tectonic
number of microorganisms present in a specific activity, which was totally unexpected on such
material. In addition, ▶ NASA and ESA have a small object. These results raised the interest of
established consensus specifications describing scientists who considered Enceladus as a prime
the number of microorganisms that are assumed objective for the Cassini mission.
to be present in a variety of materials manufactured The exploration of Enceladus by Cassini
or prepared under defined conditions. started in 2005 with a series of flybys; the closest
ones took place in March 2005 and July 2005,
with respective distances of 501 and 172 km.
See Also
Most of Enceladus’ surface is covered with
impact craters distorted by viscous relaxation
▶ Bioburden
and also by more recent tectonic activity. Cassini
▶ Microorganism
images showed several different tectonic fea-
tures: linear faults, curved stripes, and rifts
(called fossae) of over 200 km in length,
10–15 km in width, and 1 km in depth.
In addition, the Cassini instruments discov-
Enceladus ered at the South Pole a new region, previously
unobserved, different from the rest of the surface.
Therese Encrenaz This region, which extends northward up to a
LESIA, Observatoire de Paris - Section de latitude of about 55 S, is crater-free with many
Meudon, Meudon, France tectonic fractures and faults. The Cassini camera
discovered four large faults distant from each
other by about 35 km; they are up to 130 km in
Keywords length, 2 km in width, and 0.5 km in depth. They
are known as the “tiger stripes” and are the youn-
Saturn’s satellites gest structures on the satellite. The Visible and
724 Enceladus
Infrared Mapping Spectrometer of Cassini

(VIMS) detected crystalline ice in this region,
which indicates a very young origin (less than
1,000 years): if older, the water ice would have
been transformed into amorphous ice by the solar
UV radiation. The frontier between the South
Pole region and its surrounding is marked by
parallel cliffs and valleys.
After the Voyager encounters, it was already
proposed that ▶ cryovolcanism could be active on
Enceladus thus accounting for its recent
resurfacing. In addition, such cryovolcanism
would provide a source for the E-ring of ▶ Saturn,
which is located exactly at the same orbital dis-
tance as Enceladus. These hypotheses were fully
confirmed by the Cassini observations in July
2005. First, observations by the magnetometer of
Cassini showed, in the vicinity of Enceladus, a
deviation of Saturn’s magnetic field, which could Enceladus, Fig. 1 Enceladus as observed by the
be attributed to the interaction between this mag- Voyager spacecraft (#NASA)
netic field and ionized particles surrounding the
satellite. In addition, ultraviolet spectra recorded hydrocarbon chains have been identified.
during a stellar occultation by the UVIS instrument The presence of nitrogen has been interpreted as
detected a localized atmosphere, mostly made of a dissociation product of NH3, which suggests the
water vapor around the South Pole. Simulta- possible presence of liquid NH4OH associated
neously, the Cassini Ion and Neutral Mass Spec- with H2O under the surface of Enceladus.
trometer (INMS) detected not only H2O but also Enceladus’ cryovolcanism is believed to be
N2 and CO2 in the cloud surrounding the South the feeding mechanism for the E-ring of Saturn,
Pole. Finally, the Cosmic Dust Analyzer (CDA) located at the same orbital distance.
detected microcrystals of water near the tiger In June 2009, the Cosmic Dust Analyzer iden-
stripes, similar to those of the E-ring (Fig. 1). tified sodium salt (NaCl) in the icy grains that
Because of the low gravity field of Enceladus, constitute the E-ring. This constitutes a strong
the atmosphere surrounding the South Pole can- argument in favor of the presence of a salty water
not be stable and must be continuously ocean below the satellite’s surface. The presence
replenished. The most likely mechanism for feed- of salts and carbonates in the geyser products
ing this atmosphere is cryovolcanism through the suggests that a liquid ocean of water could be in
tiger stripes. This also accounts for the tempera- direct contact with a silicate core, which might
ture excess of about 15 K measured by the Com- have important implications for astrobiology.
posite Infrared Spectrometer (CIRS) around the What could be the origin of the internal energy
tiger stripes. In November 2005, a visual confir- that drives cryovolcanism on Enceladus? Tidal
mation was brought by the camera of the Cassini effects linked to the orbital resonance of
orbiter that observed jets of icy particles above Enceladus with ▶ Dione (another icy satellite of
the South Pole region. Saturn) could generate some internal energy,
Since 2005, repeated flybys have allowed sci- however, by far not enough to account for tec-
entists to accumulate data about the surface struc- tonic activity. According to some scientists,
ture of the South Pole, the geysers, and the Enceladus, as other icy satellites of Saturn,
composition of the plumes. In addition to H2O, might have formed very quickly. As a result, the
N2, CO2, and CO, other simple and complex satellite would have acquired a differentiated
Endogenicity 725
structure with a silicate core and an ice mantle.

Subsequent radioactive and tidal heating would Endergonic
have raised the inner temperature enough to melt
the icy mantle and part of the core, possibly Jacques Reisse
creating magma chambers that would feed Université Libre de Bruxelles, Brussels,
cryovolcanism. Belgium
The water torus of Enceladus has been
detected using the HIFI heterodyne spectrometer
of the ▶ Herschel space observatory, which mea- Definition
sured the water content along the line of sight and
E
a very low temperature (16 K). From estimates of An endergonic reaction is one that requires ▶ free
the external water influx rate on Saturn and Titan, energy to proceed. An example of an endergonic
it was found that the water torus of Enceladus was reaction of biological interest is ▶ photosynthe-
probably the main source of stratospheric water sis. Photosynthetic organisms conduct this reac-
for Saturn, but not for Titan. tion by using solar photons to drive the reduction
The exploration of Enceladus is the prime of carbon dioxide to glucose and the oxidation of
objective of the extended Cassini mission which water to oxygen. The free energy change during
is planned to operate until 2017. a chemical reaction or a physical transformation
which takes place at constant pressure is
See Also described by the familiar DG symbol where G is
the Gibbs free energy. By definition, DG is pos-
▶ Cryovolcanism itive for an endergonic process (and negative for
▶ Dione an ▶ exergonic process). When the reaction takes
▶ Saturn place at constant volume, the free energy change
▶ Titan is described by the symbol DF, where F is the
Helmholtz free energy.
Brown RH et al (2006) Composition and physical proper- See Also

ties of Enceladus’ surface. Science 311:1425–1428
Castillo JC et al (2006) A new understanding of the inter- ▶ Exergonic
nal evolution of Saturn’s icy satellites from Cassini
observations, 37th annual lunar and planetary science ▶ Free Energy
conference, Abstract 2200 ▶ Photosynthesis
Dougherty MK et al (2006) Identification of a dynamic
atmosphere at Enceladus with the Cassini magnetom-
eter. Science 311:1406–1409
Hansen JC et al (2006) Enceladus water vapor plume.
Science 311:1422–1425
Porco CC et al (2006) Cassini observes the south pole of
Endogenicity
Enceladus. Science 311:1393–1401
Smith BA et al (1982) A new look at the saturn system: the Nicola McLoughlin
voyager 2 images. Science 215:504–537 Department of Earth Science and Centre for
Spahn F et al (2006) Cassini dust measurements at
Geobiology, University of Bergen, Bergen,
Enceladus and implications for the origin of the
E-ring. Science 311:1416–1418 Norway
Terrile RJ, Cook AF (1981) Enceladus: “Evolution and
possible relationship with Saturn’s E-ring”, 12th annual
lunar and planetary science conference, Abstract 428
Waite JH et al (2006) Cassini ion and mass spectrometer:
Keywords
Enceladus plume composition and structure. Science
311:1419–1422 Biosignatures; Contamination
726 Endogenicity
Synonyms because they formed later in time. Often, exog-

enous components can be readily recognized
Indigenous because they occur in younger crosscutting
veins or the alteration rims of, for example,
meteorites and are derived from interactions
Definition with the terrestrial atmosphere or surface.
Demonstration of the endogenicity of a com-
Endogenicity refers to a component that maybe ponent requires its composition, structure, and
textural, chemical, mineral, or biological and that context to indicate an origin from within that
formed within the host material, for example, system. So, for instance, organic material found
a mineral phase or organic structure found in in a rock should occur in primary mineral
a rock that is indigenous to that rock and not phases or sedimentary fabrics; it should have
derived from external sources. Endogenicity con- an isotopic fingerprint that is distinct from
cerns the source of a component with respect to externally sourced, migrating organic material;
space, and this differs from ▶ syngenicity, which it should exhibit a crystallinity and/or reflec-
concerns the source of a component with respect tance that is compatible with the thermal matu-
to time. rity of the host rock. One way to demonstrate
this for carbonaceous material is to show that
the Raman spectra of the candidate biosignature
History and the host rock match, indicating that they
have experienced the same degree of thermal
Traditionally, the term endogenetic has been used alteration (e.g., Javaux et al. 2010). In addition,
in geology to refer to processes originating below carbon isotopic measurements made both in
the Earth’s surface as opposed to exogenetic, situ and on extractable phases can be used to
which describes geological processes acting at characterize and distinguish endogenetic
or near the Earth’s surface. This definition is from exogenous pools of organic material
modified here for the investigation of (e.g., Rasmussen et al. 2008). Fabric investiga-
astrobiological samples obtained from a range tions are also crucial to eliminate exogenous
of planetary surfaces and to operate at all obser- components that are hosted by younger, possi-
vational scales from the field outcrop, hand sam- bly metastable phases produced by, for exam-
ple to microscope. ple, hydrothermal alteration or weathering near
the planetary surface and occur near crosscut-
ting features or external surfaces (e.g., Pinti
Overview et al. 2009). In meteorites, for example, it has
been shown by biological staining and
It is a component formed within the host mate- epifluorescence microscopy that terrestrial
rial that is indigenous to that sample and formed microbial hyphae can rapidly colonize the inte-
in a closed system. Endogenetic components rior (Toporski and Steele 2007). Thus, storage
are not sourced from interactions with external and handling protocols are essential to protect
environments – such components are termed endogenetic remains collected by sample-return
exogenous. Most often, endogenetic compo- missions or delivered by impactors. Lastly,
nents are also syngenetic and formed at the emerging nanoscale techniques for observing
same time as the host material, but not always. the interface between a candidate biosignature
For example, diagenetic and metamorphic and host material can help to understand their
phases produced by changes in pressure and preservational history and provide a further tool
temperature after formation of the host rock for assessing endogenicity (Wacey et al. 2008;
are endogenetic to that system (provided it Thomas-Keptra et al. 2009; Bernard et al.
was closed to fluid flow) but are not syngenetic, 2007).
Endogenous Synthesis 727
See Also Definition
▶ Archean Traces of Life In geology, endogenous refers to all the processes

▶ Biogenicity that are produced in the interior of the Earth (and
▶ Biomarkers other planets). It is commonly referred to the
▶ Biomarkers, Morphological process that takes place in the ▶ mantle or the
▶ Biomarker, Isotopic core of the planets but that can have subsequent
▶ Biosignatures, Effect of Metamorphism effects on the surface of the planet. A good exam-
▶ Diagenesis ple is the mantle convection that drives the move-
▶ Raman Spectroscopy ment of tectonic plates at the surface of the Earth.
E
▶ Syngenicity The heat transfer towards its surface drives
endogenous processes. The heat derives from
the radioactive decay of elements U, Th, and
K and the residual heat from planetary accretion.
Bernard S, Benzerara K, Beyssac O, Menguy N, Guyot F,
Brown JE, Goffé B (2007) Exceptional preservation of
fossil plant spores in high-pressure metamorphic
See Also
rocks, Earth planet. Sci Lett 262:257–272
Javaux EJ, Marshall CP, Bekker A (2010) Organic-walled ▶ Exogenous
microfossils in 3.2-billion-year-old shallow-marine ▶ Geothermal Gradient
siliclastic deposits. Nature 463:934–938
▶ Mantle
Pinti DL, Mineau R, Clement V (2009) Hydrothermal alter-
ation and microfossil artefacts of the 3, 465-million- ▶ Plate Tectonics
year-old Apex chert. Nat Geosci 2:640–643
Rasmussen B, Fletcher IR, Brocks JJ, Kilburn MR
(2008) Reassessing the first appearance of eukaryotes
and cyanobacteria. Nature 455:1101–1104
Thomas-Keptra KL, Clemett SJ, McKay DS, Gibsin EK, Endogenous Synthesis
Wentworth SJ (2009) Origins of magnetite
nanocrystals in Martian Meteorite ALH84001.
Geochim Cosmochim Acta 73:6631–6677 André Brack
Toporski J, Steele A (2007) Observations from a 4-Year Centre de Biophysique Moléculaire CNRS,
contamination study of a sample depth profile through Orléans Cedex 2, France
the Martian Meteorite Nakhla. Astrobiology 7:389–401
Wacey D, Kilburn MR, McLoughlin N, Parnell J, Brasier
MD (2008) Using NanoSIMS in the search for early
life on Earth: ambient inclusion trails in a c. 3400 Ma Keywords
sandstone. J Geol Soc Lond 165:43–53
Amino acids; Formaldehyde; Hydrogen cyanide;
Hydrothermal systems; Lipids; Organic mole-
cules; Prebiotic chemistry; Primitive atmosphere
Endogenous
Daniele L. Pinti Synonyms

Geodynamics, Université du Québec à Montréal, Building blocks of primitive life
Definition
Synonyms
Life, defined as an open chemical system capable
Endogeny of self-reproduction and evolution, is thought to
728 Endogenous Synthesis
have originated from reduced organic matter in chemical bonds, respectively. The possible
water. Carbon was available as gaseous com- sources of atmospheric synthesis, including
pounds in the primitive atmosphere, either oxi- electric effects, solar UV, and impact shocks,
dized as carbon monoxide and carbon dioxide or have been reviewed by Chang (1993).
reduced as methane. When subjected to energy Miller’s laboratory synthesis of amino acids
sources (UV, heat, electric discharges, cosmic occurs efficiently when a reducing gas mixture
rays, shock waves) and mixed with water and containing significant amounts of hydrogen is
ammonia or nitrogen, these gases generate hydro- used. However, the actual composition of primi-
gen cyanide and formaldehyde that lead to many tive Earth’s atmosphere is not known. The dom-
of the building blocks of life, such as amino acids. inant view in recent years is that the primitive
Deep-sea hydrothermal systems may also repre- atmosphere consisted mainly of CO2, N2, and
sent an environment for the synthesis of prebiotic H2O along with small amounts of CO and H2
organic molecules. (Kasting and Brown 1998; Catling and Kasting
2007). Only small yields of amino acids are
formed in such a mixture (Schlesinger and Miller
Overview 1983; Miller 1998). Recent studies show that the
low yields previously reported appear to be the
It is generally believed that life on Earth arose in outcome of oxidation of the organic compounds
liquid water about 4 billion years ago. It is likely during hydrolytic workup by nitrite and nitrate
that the ingredients of primitive life were produced in the reactions. The yield of amino
organic molecules based on carbon and hydro- acids is greatly increased when oxidation inhibi-
gen atoms associated with oxygen, nitrogen, and tors, such as ferrous iron, are added prior to
sulfur. The simplest sources of carbon suscepti- hydrolysis, suggesting that endogenous synthesis
ble to lead to prebiotic organic molecules are from neutral atmospheres may be more important
gaseous, that is, carbon dioxide, carbon monox- than previously thought (Cleaves et al. 2008).
ide, and methane. A prebiotic environment must Additionally, 22 amino acids and 5 amines were
produce reduced organic molecules in which obtained when reanalyzing archived samples of
carbon is dominantly associated with hydrogen experiment run by Miller and simulating produc-
rather than oxygen. tion of organic molecules in volcanic gases by
lightning. The volcanic apparatus experiment
Production of Organics in the Atmosphere suggests that, even if the overall atmosphere
Oparin (1924) suggested that the small reduced was not reducing, localized prebiotic synthesis
organic molecules needed for primitive life were could have been effective in volcanic plumes
formed in a primitive atmosphere dominated by (Johnson et al. 2008). Unreported H2S-containing
methane. The idea was tested in the laboratory experimental samples were also found and ana-
by Miller (1953) who exposed a mixture of lyzed and were composed of numerous
methane, ammonia, hydrogen, and water to organosulfur compounds (Parker et al. 2011a,
spark discharge and silent electric discharge. In b, c). The H2S samples were analyzed by high-
his initial experiment, he obtained three amino performance liquid chromatography with fluores-
acids (glycine, alanine, and b-alanine) via the cence detection (HPLC-FD) and ultraper-
intermediary formation of hydrogen cyanide formance liquid chromatography-fluorescence
and aldehydes. More generally, simple gaseous detection with time-of-flight mass spectrometry.
molecules, like CH4, H2, NH3, and H2O, require Miller’s H2S experiment synthesized 23 amino
a supply of energy (UV, heat, electric dis- acids and 4 amines. Many nonprotein amino
charges, cosmic rays, shock waves) to react acids, including several rare or absent, in biology
with each other. They generate compounds like were detected.
formaldehyde and hydrogen cyanide that store The atmospheric formation of glycolaldehyde,
chemical energy in their double and triple a key RNA precursor, has been investigated using
a one-dimensional photochemical model. Maxi- were also run with a high-energy laser. CH4-
mum atmospheric production of glycolaldehyde containing mixtures generated hydrogen cya-
occurs when the CH4:CO2 ratio is close to 0.02. nide and acetylene, but no organics could be
The total atmospheric production rate remains obtained with CO2-rich mixtures (McKay and
small, only 1 107mol yr1. Additional produc- Borucki 1997).
tion or concentration mechanisms for
glycolaldehyde or alternative formation mecha- Submarine Hydrothermal Systems
nisms are needed (Harman et al. 2013). The reducing conditions in hydrothermal systems
The escape of hydrogen from the early Earth’s may have been an important source of biomole-
atmosphere has recently been reevaluated (Tian cules on the primitive Earth (Baross and Hoffman
E
et al. 2005). It may have occurred at rates slower 1985; Holm and Andersson 1998, 2005). The
by two orders of magnitude than previously reducing environment results from the flow of
thought. The balance between slow hydrogen substances dissolved in seawater passing inor-
escape and volcanic outgassing could have ganic compounds present in very hot crustal
maintained a hydrogen mixing ratio of more material that reduce compounds in the seawater.
than 30 %, thus making endogenous organic syn- These reduced compounds flow out of the hydro-
thesis more robust than previously thought. thermal system, and the inorganic sulfides
Intense bombardment probably caused some formed precipitate when they mix with the cold
chemical reprocessing of the Earth’s primitive (4 C) ocean water. For example, hydrocarbons
atmosphere by impact shock chemistry. An indi- containing 16–29 carbon atoms have been
cation of the number and timing of the impacts detected in the Rainbow ultramafic hydrothermal
onto the early Earth can be obtained by compar- system, Mid-Atlantic Ridge (Holm and Charlou
ison with the crater record of the Moon, which 2001). Hydrothermal vents are often disqualified
records impacts from the earliest history of the as efficient reactors for the synthesis of
solar system (Ryder 2003). Because of the larger bioorganic molecules, because of the high tem-
size of the Earth and its greater gravitational perature. However, the products that are synthe-
pull, about 20 times as many impacts would sized in hot vents are rapidly quenched in the
have occurred on the early Earth as on the surrounding cold water, which may preserve
Moon. Computer modeling of impact shock those organics formed.
chemistry shows that the nature of the atmo- In laboratory experiments, one amino acid
sphere strongly influences the shock products synthesis started with a mixture of hydrogen cya-
(Fegley et al. 1986). A neutral CO2-rich atmo- nide, formaldehyde, and ammonia, so it is not
sphere produces CO, O2, H2, and NO, while really a hydrothermal process (Hennet
a reducing CO-rich atmosphere yields primarily et al. 1992). The same holds for a reported hydro-
CO2, H2, CH4, HCN, NH3, and H2CO. The last thermal synthesis of amino acids that used cal-
three compounds are particularly interesting for cium metal, formaldehyde, ammonia hydrogen,
prebiotic chemistry since they can lead to amino and oxygen in different combinations to generate
acids via the Strecker synthesis. However, a CO- amino acids (Marshall 1994). Amino acids and
rich primitive atmosphere may be unlikely due polymers of amino acids are formed by the reac-
to the instability of CO to photolysis. In labora- tion of methane and nitrogen at 325 C in “mod-
tory experiments, a gas mixture of methane, ified seawater” in a glass-lined vessel (Yanagawa
ammonia, and water subjected to shock heating and Kobayashi 1992). “Modified seawater” con-
followed by a rapid thermal quenching yielded tains the principal metal ions present in seawater
the amino acids glycine, alanine, valine, and but at much higher concentrations so as to
leucine (Bar-Nun et al. 1970). Here again, the amplify the chemical processes catalyzed by the
gas mixture used does not represent a realistic ions so they can be more easily detected. The role
primitive atmosphere, which was probably dom- of the silicate and metal ions in this complex
inated by CO2. Laboratory simulations of shocks reaction mixture is not clear, and this system
730 Endogenous Synthesis
may not be a good model of a hydrothermal than that of Al-rich ones (montmorillonite). It is
system. assumed that every square meter of basalt or
The role of submarine hydrothermal systems komatiite rocks was punctuated by myriads of
in the synthesis of amino acids has been carefully clay-rich patches, each of them potentially
examined (Aubrey et al. 2009). The results of behaving as a single chemical reactor which
experiments exploring the potential for amino could concentrate the organics diluted in the
acid synthesis at high temperature from synthetic ocean water. Considering the high catalytic
seawater solutions of varying composition have potentiality of clays and particularly those of the
been reported. The synthesis of amino acids was Fe-rich ones (electron exchangers), it is probable
examined as a function of temperature, heating that large parts of the surface of the young Earth
time, starting material composition, and concen- participated in the synthesis of prebiotic mole-
tration. Using very favorable reactant conditions cules during the Hadean to early Archean period
(high concentrations of reactive, reduced spe- through innumerable clay-rich micro-settings in
cies), small amounts of a limited set of amino the massive parts and the altered surfaces of
acids are generated at moderate temperature con- komatiite and basaltic lavas. This leads the
ditions (125–175 C) over short heating times authors to suggest that Fe-Mg clays should be
of a few days, but even these products are signif- preferred to Al-rich ones (montmorillonite) to
icantly decomposed after exposure times of conduct experiments for the synthesis of prebi-
approximately 1 week. Although amino acids otic molecules.
can be generated from simple likely environmen- If the carbon source for life was carbon diox-
tally available precursors under submarine ide, the energy source required to reduce the
hydrothermal system conditions, the equilibrium carbon dioxide might have been provided by the
at high temperatures favors net amino acid deg- oxidative formation of pyrite from iron sulfide
radation rather than synthesis, and that synthesis and hydrogen sulfide. Pyrite has positive surface
at lower temperatures may be more favorable. charges and bonds the products of carbon dioxide
reduction, giving rise to a two-dimensional reac-
The Role of Minerals tion system, a “surface metabolism”
Clay minerals are formed by water weathering of (Wächtershäuser 1994, 1998, 2007). Laboratory
silicate minerals. As soon as liquid water was work has provided some support for this promis-
permanently present in the surface of the Earth, ing hypothesis. An early laboratory simulation of
clay minerals accumulated. The importance of hydrothermal synthetic reactions is the reduction
clay mineral in the origins of life was first of carbon dioxide to organic sulfides at 75 C in
suggested by Bernal (1949). The advantageous the presence of FeS and H2S. Methyl- and ethyl-
features of clays for Bernal were their ordered thiol were the principal thiols formed along with
arrangement, their large adsorption capacity, smaller amounts of others containing up to five
their shielding against sunlight, their ability to carbon atoms. The CO2 was also converted to
concentrate organic chemicals, and their ability CS2 and COS (Heinen and Lauwers 1996). The
to serve as polymerization templates. Since the direct reduction of CO2 to acetic acid, acetalde-
seminal hypothesis of Bernal, many prebiotic hyde, ethanol, and smaller amounts of carbon
experiments have been run with clays compounds containing up to six carbon atoms
(Ponnamperuma et al. 1982; Brack 2006; was observed to take place at 350 C and high
Negron-Mendoza et al. 2010). pressure in the presence of magnetite (FeO:
Since during the Hadean to early Archean Fe2O3) and small amounts (2 %) of water. The
period (4.5–3.5 Ga) the surface of the Earth’s yields of organics decreased 10–100 % when the
crust was predominantly composed of basalt proportion of water was increased to 90 % (Chen
and komatiite lavas, Meunier et al. (2010) con- and Bahnemann 2000). A similar inhibitory
sidered that the composition of these rocks effect of large amounts of water was observed
favored the crystallization of Fe-Mg clays rather in the reduction of nitrogen to ammonia in the
presence of magnetite and ten bars of CO2 at clay science. Elsevier Science, Amsterdam,
350 C. The yield of ammonia decreased 10- to pp 385–398
Brandes JA, Boctor NZ, Cody GD, Cooper BA, Hazen
100-fold when the water/iron ratio was increased RM, Yoder HS Jr (1998) Abiotic nitrogen reduction on
from 0.5 to 6 (Brandes et al. 1998). the early Earth. Nature 395:365–367
There are also reports of the reaction of CO in Catling D, Kasting JF (2007) Planetary atmospheres and life.
simulated hydrothermal systems. When a mixture In: Sullivan WT III, Baross JA (eds) Planets and life.
Cambridge University Press, Cambridge, pp 91–116
of CO and CH3SH was reacted with a combina- Chang S (1993) Prebiotic synthesis in planetary environ-
tion of a NiS-FeS at 100 C, acetic acid and its ments. In: Greenberg JM, Mendoza-Gomez CX,
corresponding thioester were formed (Huber and Pirronello V (eds) The chemistry of life’s origin.
Wächtershäuser 1997). This system was Kluwer Academic Publ, Dordrecht, pp 259–300
Chen QW, Bahnemann DW (2000) Reduction of carbon E
extended to the formation of keto esters at higher dioxide by magnetite: implications for the primordial
temperatures and pressures where CO is inserted synthesis of organic molecules. J Am Chem Soc
into the thioester to form a keto thioester that in 122:970–971
turn hydrolyzed to pyruvic acid (Cody et al. Cleaves HJ, Chalmers JH, Lazcano A, Miller SL, Bada JL
(2008) A reassessment of prebiotic organic synthesis
2000). More recently, a-hydroxy and a-amino in neutral planetary atmospheres. Orig Life
acids have been obtained under possible volcanic Evol Biosph 38:105–115
origin-of-life conditions by heating CO in the Cleaves HJ, Michalkova Scott A, Hill FC, Jerzy
presence of nickel or nickel/iron precipitates Leszczynski J, Nita Sahai N, Hazen R (2012)
Mineral–organic interfacial processes: potential roles
with carbonyl, cyano, and methylthio ligands as in the origins of life. Chem Soc Rev 41:5502–5525
carbon sources (Huber and Wächtershäuser Cody GD, Boctor NZ, Filley TR, Hazen RM, Scott JH,
2006). However, the plausibility of the conditions Sharma A, Yoder HS Jr (2000) Primordial
used has been debated (Bada et al. 2007). carbonylated iron-sulfur compounds and the synthesis
of pyruvate. Science 289:1337–1340
Fegley B Jr, Prinn RG, Hartman H, Watkins GH
(1986) Chemical effects of large impacts on the earth’s
primitive atmosphere. Nature 319:305–308
See Also Harman CE, Kasting JF, Wolf E (2013) Atmospheric
production of glycolaldehyde under hazy prebiotic
▶ Chemical Evolution conditions. Orig Life Evol Biosph 43:77–98
Heinen W, Lauwers AM (1996) Sulfur compounds
▶ Extraterrestrial Delivery of Organic resulting from the interaction of iron sulfide,
Compounds hydrogen sulfide and carbon dioxide in an anaerobic
aqueous environment. Orig Life Evol Biosph 26:
131–150
Hennet RJ-C, Holm NG, Engel MH (1992) Abiotic syn-
References and Further Reading thesis of amino acids under hydrothermal conditions
and the origin of life: a perpetual phenomenon?
Aubrey AD, Cleaves HJ, Bada JL (2009) The role of Naturwissenschaften 79:361–365
submarine hydrothermal systems in the synthesis of Holm NG, Andersson EM (1998) Organic molecules on
amino acids. Orig Life Evol Biosph 39:91–108 the early Earth: hydrothermal systems. In: Brack A
Bada JL, Fegley B Jr, Miller SL, Lazcano A, Cleaves HJ, (ed) The molecular origins of life: assembling pieces
Hazen RM, Chalmers J (2007) Debating evidence for of the puzzle. Cambridge University Press, Cam-
the origin of life on Earth. Science 315:937–938 bridge, pp 86–99
Bar-Nun A, Bar-Nun N, Bauer SH, Sagan C (1970) Shock Holm NG, Andersson EM (2005) Hydrothermal simula-
synthesis of amino acids in simulated primitive envi- tion experiments as a tool for studies of the origin of
ronments. Science 168:470–473 life on earth and other terrestrial planets: a review.
Baross JA, Hoffman SE (1985) Submarine hydrothermal Astrobiology 5:444–460
vents and associated gradient environment as sites for Holm NG, Charlou J-L (2001) Initial indications of abiotic
the origin and evolution of life. Orig Life Evol Biosph formation of hydrocarbons in the rainbow ultramafic
15:327–345 hydrothermal system, mid-Atlantic ridge. Earth Planet
Bernal JD (1949) The physical basis of life. Proc R Soc Sci Lett 191:1–8
Lond 357A:537–558 Huber C, Wächtershäuser G (1997) Activated acetic acid
Brack A (2006) Clay minerals and the origin of life. In: by carbon fixation on (Fe, Ni)S under primordial con-
Bergaya F, Theng BKG, Lagaly G (eds) Handbook of ditions. Science 276:245–247
732 Endogeny
Huber C, Wächtershäuser G (2006) a-hydroxy and Wächtershäuser G (1994) Life in a ligand sphere. Proc
a-amino acids under possible Hadean, volcanic origin- Natl Acad Sci U S A 91:4283–4287
of-life conditions. Science 314:630–632 Wächtershäuser G (1998) Origin of life in an iron-sulfur
Johnson AP, Cleaves HJ, Dworkin JP, Glavin DP, world. In: Brack A (ed) The molecular origins of life:
Lazcano A, Bada JL (2008) The Miller volcanic assembling pieces of the puzzle. Cambridge Univer-
spark discharge experiment. Science 322:404 sity Press, Cambridge, pp 206–218
Kasting JF, Brown LL (1998) The early atmosphere as a Wächtershäuser G (2007) On the chemistry and
source of biogenic compounds. In: Brack A (ed) The evolution of the pioneer organism. Chem Biodivers
molecular origins of life: assembling pieces of the 4:584–602
puzzle. Cambridge University Press, Cambridge, Yanagawa H, Kobayashi K (1992) An experimental
pp 35–56 approach to chemical evolution in submarine
Marshall WL (1994) Hydrothermal synthesis of amino hydrothermal systems. Orig Life Evol Biosph 22:
acids. Geochim Cosmochim Acta 58:2099–2106 147–159
McKay CP, Borucki WJ (1997) Organic synthesis in
experimental impact shocks. Science 276:390–392
Meunier A, Petit S, Cockell CS, El Albani A, Beaufort
D (2010) The Fe-rich clay microsystems in basalt-
komatiite lavas: importance of Fe-smectites for Endogeny
pre-biotic molecule catalysis during the Hadean Eon.
Orig Life Evol Biosph 40:253–272 ▶ Endogenous
Miller SL (1953) The production of amino acids under
possible primitive Earth conditions. Science
117:528–529
Miller SL (1998) The endogenous synthesis of organic
compounds. In: Brack A (ed) The molecular origins Endolithic
of life: assembling pieces of the puzzle. Cambridge
University Press, Cambridge, pp 59–85
Negron-Mendoza A, Ramos-Bernal S, Mosqueira FG Angeles Aguilera
(2010) The role of clay interactions in chemical evo- Laboratorio de Extremófilos, Centro de
lution. In: Basiuk VA (ed) Astrobiology: emergence, Astrobiologı́a (INTA-CSIC), Torrejón de Ardoz,
search and detection of life. American Scientific Pub-
lishers, Stevenson Ranch, pp 214–233
Madrid, Spain
Oparin AI (1924) Proikhozndenie Zhizni. Izd. Moskowski
Rabochi, Moscow
Parker ET, Cleaves HJ, Callahan MP, Dworkin JP, Definition
Glavin DP, Lazcano A, Bada JL (2011a) Prebiotic syn-
thesis of methionine and other sulfur-containing organic
compounds on the primitive Earth: a contemporary Endolithic refers to any ▶ microorganisms
reassessment based on an unpublished 1958 Stanley which are able to colonize and survive inside
Miller experiment. Orig Life Evol Biosph 41:201–212 a rock. Endoliths are classified into three differ-
Parker ET, Cleaves HJ, Dworkin JP, Glavin DP,
Callahan M, Aubrey A, Lazcano A, Bada JL (2011b)
ent classes depending on the part of the interior
Primordial synthesis of amines and amino acids in a of the rock that the organisms colonize:
1958 Miller H2S-rich spark discharge experiment. chasmoendoliths (chasm means cleft) are the
Proc Natl Acad Sci U S A 108:5526–5531 organisms that colonize open spaces in the rock
Parker ET, Cleaves HJ, Callahan MP, Dworkin JP, Glavin
DP, Lazcano A, Bada JL (2011c) Enhanced synthesis
such as fissures and cracks; cryptoendoliths
of alkyl amino acids in Miller’s 1958 H2S experiment. (crypto means hidden) are the organisms that
Orig Life Evol Biosph 41:569–574 colonize the interior of porous rocks; and
Ponnamperuma C, Shimoyama A, Friebele E (1982) Clay euendoliths (eu which means true endolith) are
and the origin of life. Orig Life 12:9–40
Ryder G (2003) Bombardment of the Hadean Earth:
the organisms that actively colonize the interior
wholesome or deleterious? Astrobiology 3:3–6 of the rock by forming tunnels with the shape of
Schlesinger G, Miller SL (1983) Prebiotic syntheses in their bodies. These organisms are of extreme
atmospheres containing CH4, CO, and CO2 1. Amino astrobiological interest because the interior of a
acids. J Mol Evol 19:376–382
Tian F, Toon OB, Pavlov AA, De Sterck H (2005)
▶ meteorite can give shelter to endolithic organ-
A hydrogen-rich early atmosphere. Science isms under the extreme conditions of an
308:1014–1017 interplanetary voyage (▶ lithopanspermia).
Endosymbiosis 733
See Also Definition
▶ Cryptoendolithic Endosymbiosis is a symbiotic association in

▶ Lithopanspermia which one partner, generally a prokaryote sym-
▶ Microorganism biont, lives sequestered inside specialized
▶ Panspermia eukaryotic host cells called bacteriocytes. The
association is obligate for both partners.
Endospore
Overview E
1,2 3
Wayne L. Nicholson and Ralf Moeller
1 Symbiogenesis, the evolutionary process of
Space Life Sciences Laboratory, University of
establishing a symbiotic association, has been
2 one of the dominant forces in the early evolution
Space Life Sciences Laboratory, Kennedy Space
of life on Earth. In 1967 Lynn Margulis postu-
Center, University of Florida, Gainesville, FL, USA
3 lated the serial endosymbiotic theory of eukary-
otic ▶ cell evolution, currently accepted with
respect to the origin of mitochondria and ▶ chlo-
roplasts (Margulis 1993). These two eukaryotic
Synonyms ▶ organelles are beyond doubt the product of
symbiogenic events between prokaryotes and
Bacterial spore; Spore primitive eukaryotes, as supported by many dif-
ferent types of genetic, biochemical, and phylo-
genetic evidence. The origin of mitochondria
Definition dates back to 2 109 years ago, whereas chloro-
plasts originated more than 1.2 109 years ago,
An endospore is a tough, dormant structure produced because fossil red algae of that age have been
by certain species of Gram positive bacteria, most found. DNA sequence comparisons and phyloge-
notably of the genera Bacillus and Clostridium. netic analyses have identified alpha-
proteobacteria and cyanobacteria as the bacterial
groups to which the ancestors of mitochondria
See Also
and plastids, respectively, belonged.
Nowadays, symbiotic associations have been
▶ Sporulation
documented in practically every major branch of
the ▶ tree of life, and this observation reinforces
the role played by ▶ symbiosis in the emergence
Endosymbiosis of evolutionary innovations in eukaryotes.
Recently, the genomic era has opened the access
Amparo Latorre to the characterization of the organisms involved
Institute Cavanilles for Biodiversity and in symbiosis, mainly those non-cultivable micro-
Evolutionary Biology, Universitat de Valencia, bial endosymbionts, allowing the comparison
Valencia, Spain among the different evolutionary innovations
carried out by these bacteria on their way from
a free-living lifestyle to varied stages of integra-
Keywords tion with their respective hosts (Dale and Moran
2006; Moya et al. 2008; McCutcheon and Moran
Bacteriocyte; Chloroplast; Endosymbiont; Mito- 2012). Thus, endosymbiosis is an ongoing phe-
chondria; Obligate mutualism; Symbiogenesis nomenon in the evolution of life and played a key
734 Endothermic
role in the origin and evolution of the McCutcheon JP, Moran NA (2012). Extreme genome
eukaryotic cell. reduction in symbiotic bacteria. Nat Rev Microbiol
10:13–26.
In general, it is well established that endosym- McCutcheon JP, von Dohlen CD (2011) An
biotic integration is a process that commonly interdependent metabolic patchwork in the nested
changes profoundly the gene repertoire of the symbiosis of mealybugs. Curr. Biol. 21:1366–72.
free-living ancestor. Depending on the type of Moran NA et al (2008) Genomics and evolution of heri-
table bacterial symbionts. Annu Rev Genet
symbiotic relationship (i.e., mutualistic or para- 42:165–190
sitic, facultative or obligate, etc.), age of the Moya A et al (2008) Learning how to live together: geno-
association, and host necessities (i.e., nutritional, mic insights into prokaryote-animal symbioses. Nat
defensive, waste recycling, etc.), the genetic and Rev Genet 9:218–229
Nakabachi A et al (2006) The 160-kilobase genome of
metabolic changes will be more or less dramatic the bacterial endosymbiont Carsonella. Science 314:
(Wernegreen 2005; Moran et al. 2008). 267
We should not disregard the discovery of new Nowack ECM et al (2008) Chromatophore genome
instances of endosymbionts currently en route to sequence of Paulinella sheds light on acquisition
of photosynthesis by eukaryotes. Curr Biol 18:
become organelles or fully fledged eukaryotic 410–418
compartments of bacterial origin (Nakabachi Pérez-Brocal V et al (2006) A small microbial genome:
et al. 2006; Pérez-Brocal et al. 2006; Nowack the end of a long symbiotic relationship? Science
et al. 2008; McCutcheon and von Dohlen 2011). 314:312–313
Wernegreen JJ (2005) For better or worse: genomic con-
Current “omics” (such as genomics, proteomics, sequences of intracellular mutualism and parasitism.
or metabolomics) methodologies allow us not Curr Opin Genet Dev 15:572–583
only to further unravel the steps toward the origin
of the eukaryotic cell but also to assess the ques-
tion of how much of the eukaryotic complexity
originated through evolutionary innovations by
symbiogenesis. Endothermic
Jacques Reisse
Université Libre de Bruxelles, Brussels,
See Also
Belgium
▶ Cell
▶ Chloroplast
Definition
▶ Eukarya
An endothermic reaction is a physical transfor-
▶ Mitochondrion
mation which requires an energy supply if it takes
▶ Organelle
place at constant volume or which requires an
▶ Phylogenetic Tree
enthalpy supply if it takes place at constant pres-
▶ Phylogeny
sure. A familiar endothermic physical transfor-
▶ Symbiosis
mation is the evaporation of water. The enthalpy
▶ Taxonomy
change (DH) or the energy change (DU) during an
endothermic reaction or transformation is posi-
References and Further Reading tive by definition.
Dale C, Moran NA (2006) Molecular interactions

between bacterial symbionts and their host. Cell
126:453–465
See Also
Margulis L (1993) Symbiosis in cell evolution, 2nd edn.
W.H. Freeman, San Francisco, p 452 ▶ Exothermic
Energy 735
Overview
Energy
Ever since the Industrial Revolution, one of man-
Francisco Montero kind’s greatest challenges has been to find inter-
Department of Biochemistry and Molecular conversion mechanisms for different energy forms
Biology I, Facultad de Ciencias Quı́micas, (i.e. conversion of thermal, wind, electromagnetic,
Universidad Complutense de Madrid, Madrid, or atomic energy into mechanical or electrical
Spain energy). Although all energy forms are intercon-
vertible, there are limitations with regard to the
amount and quality of the energy released in a
E
Keywords process that can be used in another one. For this
reason, the total energy in a system is usually
ATP; Bioenergetics; Chemical work; Electro- classified into different categories, such as internal
chemical potential gradient; Electromagnetic energy, free energy, enthalpy, entropic contribu-
energy; Energy transduction; Osmotic work tion, etc. Of these, free energy is the only one that
can be transferred from one process to another.
Therefore, processes are classified as ▶ ender-
Definition gonic or ▶ exergonic depending, respectively, on
whether they need or release ▶ free energy during
In physics, energy is defined as the capacity of a their development.
system to produce work. There are many forms of Biological systems, when viewed as thermody-
energy (e.g., mechanical, electrical, electromag- namic systems, are considered to be open systems,
netic, chemical, etc.), and it is possible to convert which means they can exchange matter and energy
some energy forms into others, with the condition with their surroundings in order to maintain the
that the energy is conserved. dissipative structures, which characterize them.
Basically, energy can be exchanged with the sur-
roundings in two ways: through matter, by taking
History molecules from the environment whose catabolic
processes (of degradation) release free energy, or
A coherent theory on energy and its transforma- by absorbing electromagnetic radiation. Further-
tions was first advanced in the nineteenth century, more, biological systems have mechanisms which
chiefly as a consequence of the Industrial Revo- can convert some forms of energy into others, e.g.,
lution. The propounding of the fundamental prin- electromagnetic energy into osmotic energy,
ciples (or laws) of thermodynamics established osmotic energy into chemical and mechanical
the basic rules for energy conversions. energy, etc. These general mechanisms for the
Although important observations had already translation of energy within a biological system,
been made on the effect of energy on biological which are the result of evolution and natural selec-
systems by the nineteenth century, it was not until tion, constitute the field of study of bioenergetics
the development of biochemistry in the twentieth and are truly astonishing. A general law governing
century that energy conversion mechanisms in bio- energy coupling in biological systems states that
logical systems were explained. In 1941, Lipmann they are not usually direct. Instead, they are pro-
established that ATP is an energy currency in the duced through universal intermediaries or “energy
biological system, while in 1961, Mitchell pro- currencies.” An exergonic process produces an
posed the chemiosmotic hypothesis, which consti- energy currency which can then be used in an
tutes the base of all modern ▶ bioenergetics. endergonic process. All of the energy currencies
Nowadays, the majority of energy transduction can be used to produce chemical work (production
processes at molecular level are well understood. of molecules whose formation requires energy, in
736 Energy Conservation
general anabolic processes), osmotic work (the converted from one form to another; however,
transport of substances across membranes against the law of conservation of energy, a fundamental
their ▶ electrochemical potential gradients), or law of physics, states that although energy can be
mechanical work (movement of flagella and cilia, changed in form, it can be neither created nor
contractile movements, etc.). destroyed. The main energy conservation mech-
anisms used by biological systems are ▶ respira-
tion (aerobic and anaerobic), fermentation, and
See Also
▶ photosynthesis (oxygenic and anoxygenic).
▶ ATP
▶ ATP Synthase See Also
▶ Bioenergetics
▶ Electrochemical Potential ▶ Aerobic Respiration
▶ Endergonic ▶ Anaerobic Respiration
▶ Endothermic ▶ Anoxygenic Photosynthesis
▶ Energy Conservation ▶ ATP
▶ Energy Sources ▶ ATPase
▶ Enthalpy ▶ ATP Synthase
▶ Entropy ▶ Bioenergetics
▶ Exergonic ▶ Electron Acceptor
▶ Exothermic ▶ Electron Carrier
▶ Free Energy ▶ Electron Transport
▶ Energy
▶ Energy Sources
▶ Fermentation
Nichols DG, Ferguson SJ (2013) Bioenergetics4. Aca- ▶ NADH, NADPH
demic Press, London ▶ Oxidation
Skulachev VP (1992) The laws of cell energetics. Eur ▶ Photosynthesis
J Biochem 208:203–209 ▶ Photosynthesis, Oxygenic
Skulachev VP (1994) Bioenergetics: the evolution of molec-
ular mechanisms and the development of bioenergetics ▶ Proton Motive Force
concepts. Antonie Van Leeuwenhoek 65:271–284 ▶ Proton Pump
▶ Respiration
Energy Conservation
Energy Sources
Juli Peretó
Institut Cavanilles de Biodiversitat i Biologia Felipe Gomez
Evolutiva, Universitat de València, València, Spain Centro de Astrobiologı́a (CSIC/INTA), Instituto
Definition
▶ Energy conservation refers to the mechanisms Keywords

by which organisms can transform different
▶ energy sources (radiation, reduced organic or Electron donors; Energy conservation; Fermen-
inorganic compounds) into cellular useful tation; Photosynthesis; Radiation; Redox reac-
energy: proton motive force, ▶ ATP, and reductions; Reduced inorganic compounds; Reduced
ing power. It is common for energy to be organic compounds; Respiration
Energy Sources 737
Definition compounds like ▶ ATP or NADH. Some reac-

tions or functions are driven by the proton
Life implies work, in other words, it requires motive force (transport of metabolites through
▶ energy. Metabolite transport across mem- the membrane, the rotation of the bacterial fla-
branes, cellular ▶ motility, or the biosynthesis gella); others require the use of ATP, such as the
of macromolecules are examples of cellular pro- synthesis of macromolecules. An important
cesses that require energy. Any living system achievement was the establishment of the
relies on an external source of energy. The exter- chemiosmotic hypothesis coupling the ▶ elec-
nal sources of energy useful for living systems are tron transport in the respiratory chain with the
only two: radiation and reduced chemical com- generation of a proton motive force and the syn-
E
pounds. Living systems transform external thesis of ATP proposed by Peter Mitchell in
energy sources into energy useful for cellular 1961, and for which he was awarded the Nobel
processes according to the thermodynamic law Price.
of energy conservation.
See Also
History
▶ ATP Synthase
Peter Mitchell was awarded the 1978 Nobel Prize ▶ Chemotroph
in Chemistry for his work in the description of the ▶ Electrochemical Potential
Chemiosmosis. Paul D. Boyer and John ▶ Electron Donor
E. Walker were awarded part of the 1997 Nobel ▶ Electron Transport
Prize in Chemistry for their clarification of the ▶ Energy
mechanism of ATP synthesis. ▶ Fermentation
▶ Metabolism
▶ Motility
Overview ▶ Photosynthesis
Energetic processes are basically related to ▶ Radiation
oxido-reduction reactions associated with the ▶ Respiration
transformation of the energy sources. Energy
sources are key elements for the classification
of organisms. According to this, organisms can
be classified as chemotrophs if the external Amend JP, Shock EL (2001) Energetics of overall meta-
source of energy is chemical or phototrophs if bolic reactions of thermophilic and hyperthermophilic
it is radiation. Chemotrophs can be subdivided archaea and bacteria. FEMS Microbiol Rev
as chemoorganotrophs if the source of energy is 25:175–243
Mitchell P (1967) Proton current flow in mitochondrial
a reduced organic compound (e.g., glucose, ace- systems. Nature 25(5095):1327–1328
tate), or chemolithotrophs if the source of energy Mitchell P, Moyle J (1967) Chemiosmotic hypothesis
is a reduced inorganic compound (e.g., ferrous of oxidative phosphorylation. Nature 213(5072):
iron, ammonia). The mechanisms by which 137–139
Smith E, Morowitz HJ (2004) Universality in intermedi-
organisms transform the external source of ary metabolism. Proc Natl Acad Sci U S A
energy into cellular energy are three: ▶ respira- 101:13168–13173
tion, ▶ fermentation, which is used by Temple KL, Colmer AR (1951) The autotrophic oxidation
chemothrophs, and ▶ photosynthesis, which is of iron by a new bacterium: thiobacillus ferrooxidans.
J Bacteriol 62(5):605–611
used by phototrophs. Cells can store the Wächtershäuser G (1992) Groundworks for an evolution-
transformed energy as ▶ proton motive force ary biochemistry - the iron sulphur world. Prog
(proton gradient) or in the form of energetic Biophys Mol Biol 58:85–201
738 Enthalpy
of life. For example, the semi-phosphorylative

Enthalpy ED pathway has been demonstrated in several
anaerobic bacteria and in the halophilic archaea,
Jacques Reisse whereas the non-phosphorylative ED pathway is
Université Libre de Bruxelles, Brussels, Belgium found in some fungi and thermophilic and hyper-
thermophilic archaea. It has been suggested that
the ED pathway is older than the EMP pathway.
Definition
The enthalpy change is the heat produced or History

absorbed during a chemical or physical transfor-
mation taking place at constant pressure. The The ED pathway was first discovered in 1952 in
thermodynamic definition of enthalpy is H = Pseudomonas saccharophila and in 1967 was
U + PV, where H stands for enthalpy, U for shown to be also present in Escherichia coli.
internal energy, P for pressure, and V for volume.
See Also
See Also ▶ Embden-Meyerhof-Parnas Pathway

▶ Metabolism
▶ Free Energy
Entropy
Entner-Doudoroff Pathway
Jacques Reisse
Juli Peretó Université Libre de Bruxelles, Brussels, Belgium
Spain Definition
Entropy is a thermodynamic concept introduced

Synonyms in 1865 by Clausius. Entropy, represented by S,
increases in any spontaneous processes (where
Glycolysis DS is positive) taking place in an isolated system,
i.e., a system which does not exchange matter or
energy with its surroundings. If two gases are in
Definition a constant-temperature box, initially separated by
a partition and that partition is removed, the gases
The Entner-Doudoroff (ED) pathway is the gly- will mix without a change in enthalpy. Entropy,
colytic pathway that allows to catabolize sugar however, increases as the gases mix in this spon-
acids, such as gluconate. Several enzymatic steps taneous process and acts as the driving force.
allow the conversion of six carbon sugar acids in
two trioses that can be further metabolized
through the universal lower part of the See Also
▶ Embden-Meyerhof-Parnas (EMP) pathway.
This pathway is present in bacteria, but modifi- ▶ Enthalpy
cations have been described in all three domains ▶ Free Energy
Environment 739
ecological niche and the activities that take

Environment place within it. Culture-independent molecular
biology techniques, such as in situ hybridization
José Luis Sanz with fluorescent probes (FISH) or those based
Departamento de Biologı́a Molecular, on PCR- denaturing gradient gel electrophoresis
Universidad Autónoma de Madrid, Madrid, or the creation of genetic libraries, have given
Spain rise to “molecular microbial ecology” allowing
great steps to be made in the understanding
of microbial communities that live in specific
Keywords habitats that would have been impossible with
E
traditional microbiological research techniques.
Acidity; Atmosphere; Ionic strength; Life; Microorganisms occupy extraordinarily
Molecular ecology; Oceans; Pressure; Radiation; diverse habitats needing only liquid water to
Redox conditions; Temperature develop. They not only thrive in soils and water
(fresh or marine) with “normal” physical/chemi-
cal conditions but have also adapted to ▶ extreme
Synonyms environments: low and high temperatures and
pHs, high salt concentrations, high pressure,
Ecosystem; Habitat etc. Furthermore, given the diminutive size of
microorganisms, microniches or microenviron-
ments can exist in conditions, which may differ
Definition greatly from those of the general environment,
where they are contained. This is especially
Natural environment refers to the collection important in sediments and subsurfaces where
of objects both living and inanimate as well as the semisolid structure of the substrate allows
the fluid, gaseous, or solid media that together microniches, whose properties may vary dramat-
make up a ▶ habitat. The term includes soil, ically to coexist in proximity. Environmental
the subsurface, the atmosphere, the oceans and conditions are very important in astrobiology
smaller bodies of water, and the interiors because they can determine the habitability of a
of plants and animals. The description of an given planet or planetary body.
environment requires definitions of its physical
and chemical characteristics – temperature
and pressure, acidity, redox conditions, See Also
radiation, etc.
▶ Biosphere
Overview ▶ Biotope
▶ Ecosystem
The ensemble of biological communities and ▶ Extreme Environment
their surroundings constitutes an “▶ ecosys- ▶ pH
tem,” and the science that studies these ecosys-
tems is known as Ecology. From a
microbiological point of view, “environmental
microbiology” studies the microbial processes References and Further Reading
that take place in the soil, water, food,
Johnson DL, Ambrose SH, Bassett TJ, Bowen ML,
depuration systems, etc., and “microbial ecol-
Crummey DE, Isaacson JS, Johnson DN, Lamb P,
ogy” describes qualitatively and quantitatively Saul M, Winter-Nelson AE (1997) Meanings of envi-
the ▶ microorganisms that occupy a specific ronmental terms. J Environ Qual 26:581–589
740 Environmental Genome
Madigan M, Martinko J, Dunlap P, Clark D (2009) Brock Definition

biology of microorganisms, 12th edn. Person Educa-
tion/Benjamin Cummings, San Francisco
Schlesinger WH (2005) Biogeochemistry. In: Schlesinger An enzyme is a biological ▶ catalyst necessary
WH (ed) Treatise on geochemistry, vol 8. Elsevier for metabolic reactions to occur at an adequate
Science, Amsterdam rate under mild conditions of pH and tempera-
ture, typically consisting largely or entirely of
protein, though catalytic RNA molecules are
also known.
Environmental Genome
History
▶ Metagenome
Enzyme catalysis was discovered by Lazzaro
Spallanzani in the eighteenth century as an
observation that the digestion of food in the
Environmental Sequence animal stomach was chemical rather than
a purely mechanical process of grinding. Subse-
▶ Phylotype quent advances in the nineteenth century came
primarily from studies of digestion and of the
action of yeast in fermentation. Until the early
twentieth century, the usual word for enzyme
was ferment, now entirely superseded by
Environmental Transcriptome
enzyme, a name that also referred originally to
yeast, as it was coined by Wilhelm K€uhne from
▶ Metatranscriptome
Greek words meaning “in yeast.” When the last
printed version of Enzyme Nomenclature was
published in 1992, about 3,200 distinct enzyme
activities were recognized, but the current num-
Enzyme ber is about 6,500, and the number of distinct
proteins known to be enzymes is very much
Athel Cornish-Bowden and Marı́a Luz Cárdenas larger.
Unité de Bioénergétique et Ingénierie des
Protéines, Centre National de la Recherche
Scientifique, Aix-Marseille Université, Marseille Overview
Cedex 20, France
All metabolic reactions are catalyzed by specific
molecules called enzymes. Even reactions that
Keywords proceed spontaneously and rapidly, such as the
hydration of CO2, are normally catalyzed in
Active site; Allosteric site; Catalysis; Isoenzyme; physiological systems. Before the discovery of
Kinetics; Metabolism; Ribozyme catalytic RNA molecules (sometimes called
ribozymes), all enzymes were believed to con-
sist mainly or entirely of protein molecules, typ-
Synonyms ically with more than 100 ▶ amino acid residues
in each subunit. It is now widely accepted that
Ferment (obsolete) catalytic RNA satisfies the definition of an
Enzyme 741
enzyme, and the ▶ ribosome is a fundamental to respond with high sensitivity to changes in
example of catalytic RNA (Steitz and Moore conditions. The part of an enzyme where cataly-
2003). Biological catalysts include metabolites sis occurs is called the active site, and if a differ-
that participate in cycles of reactions, such as ent site exists for binding effectors, it is called an
citrate, regenerated in each turn of the allosteric site.
tricarboxylate (Krebs) cycle, but also many Enzymes are grouped in Enzyme Nomenclature
other metabolites. These are certainly catalysts, into six classes, of which three (oxidoreductases,
but they are not conventionally regarded as transferases, and hydrolases) catalyze group-
enzymes; however, from the point of view of transfer reactions and three (lyases, isomerases,
astrobiology, they probably should be regarded and ligases) catalyze other types of reaction. This
E
as enzymes because at the origin of life, there is a classification of reactions catalyzed, not a clas-
must have been catalysts produced by the first sification of proteins catalyzing them. The number
organisms that were neither protein nor RNA, of known proteins acting as enzymes is very much
but much simpler molecules. larger than the number of known activities
Although the effectiveness of enzymes as (because the proteins catalyzing any given reaction
efficient catalysts is normally emphasized, in different organisms are nearly always different),
their fundamental property is specificity because and the amount of variation in structure among the
without specificity, there could be no organiza- known cases suggests that the total number is
tion or regulation of metabolic processes. Most huge. In most of the known cases, the proteins
enzymes catalyze just one or a small number of are similar enough to be clearly homologous, but
reactions and are largely or completely inactive some exceptions, exemplified by superoxide
in others that appear quite similar. Bacterial dismutase, are known. Even within a single spe-
glucokinase, for example, catalyzes the phos- cies, it is common to find multiple proteins, known
phorylation of glucose by ATP, but does not as isoenzymes, catalyzing the same reaction, often
catalyze the phosphorylation of a hexose as with distinct regulatory properties.
similar as mannose. Even relatively unspecific
enzymes, mostly involved in catabolic pro-
cesses such as digestion, are highly specific by
See Also
the standards of inorganic catalysts. Another
property exhibited by all enzymes is saturation:
▶ Amino Acid
although the rate of reaction may be propor-
▶ Metabolism
tional to the concentration of each reactant at
low concentrations, it never increases indefinitely
▶ Ribosome
with increases in concentration, the rate
▶ Ribozyme
v typically depending on the substrate concentra-
tion a according to the Michaelis-Menten equa-
tion, v = Va/(Km + a), in which V, the limiting References and Further Reading
rate, and Km, the Michaelis constant, are con-
stants. In vivo, however, enzymes typically oper- Cornish-Bowden A (2012) Fundamentals of enzyme
kinetics. Wiley-Blackwell, Weinheim
ate in or close to the proportional zone, in other
Enzyme Nomenclature. http://www.chem.qmul.ac.uk/
words at substrate concentrations similar to or iubmb/enzyme/
smaller than Km. Certain enzymes, known as Fersht AR (1998) Structure and mechanism in protein
regulatory enzymes, are inhibited or activated science: a guide to enzyme catalysis and protein fold-
ing. Freeman, San Francisco
by molecules structurally different from their
Steitz TA, Moore PB (2003) RNA, the first macromolec-
substrates or products, which are called effectors, ular catalyst: the ribosome is a ribozyme. Trends
or display special kinetic laws that enable them Biochem Sci 28:411–418
742 Enzymology: History of
Enzymology: History of Ephemeris
Stéphane Tirard Daniel Rouan

Centre François Viéte d’Histoire des Sciences et LESIA, Observatoire Paris-Site de Meudon,
des Techniques EA 1161, Faculté des Sciences et Meudon, France
Definition
Definition
An ephemeris is the set of tabulated coordinates
In 1833, the French biologists A. Payen of an astronomical object in the sky at successive
(1795–1871) and J. F. Persoz (1805–1868) iso- dates. Plural: ephemerides.
lated a malt-soluble ferment able to digest the
amide and called it diastase. In 1878, the German
physiologist Wilhelm K€uhne (1837–1900)
named the contents of this digestive juice as See Also
enzyme. During the second part of the nineteenth
century, there was an active debate between ▶ Coordinate Systems
Louis Pasteur (1822–1895) and Justus Freiherr ▶ Declination
von Liebig (1803–1873) about the cause of fer- ▶ Right Ascension
mentation. Pasteur argued that fermentation is a
process of microscopic living entities, like yeast,
and Liebig argued that fermentation is a sponta-
neous decomposition of matter.
In 1897, the German chemistry Ed€uard Epilithic
Buchner (1860–1917) made an in vitro fermen-
tation of sugars in an acellular fraction of yeast. Gerda Horneck
Some authors said that this result probably con- DLR German Aerospace Center, Institute of
stituted the beginning of biochemistry science. Aerospace Medicine, Radiation Biology, Köln,
A few years later, in 1903, the French-Russian Germany
physical chemist Victor Henry (1872–1940)
stated that all enzymes are proteins.
The 1920s have witnessed important develop-
ments of enzymology with the purification and Definition
the crystallization of numerous enzymes (urease,
pepsin, trypsin, etc.) and the study of the enzy- Epiliths are organisms or microbial communities
matic kinetic. that live on the surface of rocks. Examples are
▶ lichens, mosses, biofilms, and desert varnish.
Epiliths are generally well adapted to resist envi-
ronmental extreme conditions, such as large var-
See Also iations in temperature, extreme dryness, and
intense solar irradiation. They are sometimes
▶ Catalyst the only colonizers in desert and high-mountain
▶ Enzyme regions.
Equation of State 743
See Also undertaken (http://epoxi.umd.edu/1mission/sta-

tus.shtml).
▶ Biofilm The team working with EPOXI lost commu-
▶ Desiccation nication to the spacecraft sometime between
▶ Endolithic August 11 and 14, 2013. On Monday, September
▶ Extreme Environment 16, 2013, after considerable effort transmitting
▶ Extreme Ultraviolet Light low-level hardware commands, EPOXI team
▶ Hypolithic determined that there were no other plausible
▶ Lichens scenarios under which they could recover com-
▶ Microbial Mats mand and control of the spacecraft. They
E
recommended that NASA declares the mission
lost. That declaration was announced by NASA
on Thursday, September 19, 2013.
Episome
▶ Plasmid
See Also
▶ Deep Impact
▶ Transiting Planets
EPOXI Mission
References
William M. Irvine
University of Massachusetts, Amherst, MA, USA A’Hearn MF et al (2013) EPOXI at Comet Hartley 2. Sci-
ence 332:1396–1400
Definition
EPOXI was an extension of the NASA Deep

EPS
Impact mission (▶ Deep Impact) with two
▶ Exopolymers
goals: to observe extrasolar planets using the
transit method and to study comet 103P/Hartley
2 using the same instruments used for the Deep
Impact observations of comet 9P/Tempel 1. Sev-
eral transiting planets were observed, and Equation of State
Earth was also observed, in an effort to see
what characteristics of Earth-like planets might Tristan Guillot
be most easily measured remotely. The closest Observatoire de la Côte d’Azur, Université de
approach to comet 103P/Hartley 2 occurred on Nice-Sophia Antipolis, CNRS, Nice, France
November 4, 2010. The name EPOXI is a hybrid
of EPOCH (Extrasolar Planet Observation and
CHaracterization Investigation) and DIXI (Deep Keywords
Impact eXtended Investigation). The EPOXI
Mission web page maintained by NASA Density; Pressure; Temperature;
includes status reports on the various activities Thermodynamics
744 Equinox
Definition They describe how matter is affected by the

extreme conditions that occur there. With a pres-
An equation of state is a relation (or by extension sure of about 360 GPa (about 3.6 million times
a series of relations) between thermodynamic state the atmospheric pressure at sea level on Earth)
variables characterizing matter in a given state, for and a temperature of about 7,000 K, the iron
instance, in a celestial body. It describes how a mac- which forms most of the core at the center of
roscopic entity is affected by external changes (e.g., the Earth is believed to be in solid form. How-
changes in temperature, pressure, volume, etc.). ever, the equation of state of iron tells us that at
When modeling stars and planets, equations of conditions relevant slightly higher up in the
state are used in particular to calculate density as Earth, it should be liquid, therefore forming the
a function of pressure and temperature, the phases so-called liquid outer core. In giant planets such
(e.g., gaseous, liquid, solid, etc.) of the materials as Jupiter and Saturn, the equation of state of
considered, and how temperature changes with hydrogen indicates that this element ionizes
pressure during macroscopic motions. from a molecular state, H2, to a conducting
state called metallic hydrogen at pressures of
about 150 GPa and temperatures of 10,000 K
Overview or less.
The first and most widely known equation of state

is the so-called ideal gas law. It links the pressure See Also
P, volume V, absolute temperature T, and number
of particles N in a gas composed of nonrelativistic, ▶ Atmosphere, Structure
noninteracting mass particles through the relation ▶ Interior Structure, Planetary
PV = NkT, where k = 1.38065034 1023 J K1 is ▶ Low Mass Star
Boltzmann’s constant. Equations of state of arbi- ▶ Stellar Evolution
trary complexity can be calculated for elements
that undergo a phase transition (i.e., become gas-
eous, liquid, or solid), for mixtures, for photons,
for charged particles, in the presence of magnetic
Equinox
fields, etc. Among other things, equations of state
are crucial to model atmospheric structures: when
Daniel Rouan
a parcel of air (or fluid) is transformed (e.g., by its
motion) without losing heat (i.e., adiabatically), it
Meudon, France
conserves its ▶ entropy. Entropy is a thermody-
namic state variable and, as such, it also obeys an
equation of state relation that links it to the other
thermodynamic state variables of the problem Synonyms
(e.g., temperature, pressure). Using its conserva-
tion, one can therefore calculate the rate of change Autumnal point; Vernal point
of temperature with pressure in such a transforma-
tion. In atmospheres that are thick enough for
convection to occur nearly adiabatically, the tem- Definition
perature profile is mostly set by the temperature of
the photosphere (the level at which most of the The equinox is one or the other of the two points on
photons can escape freely to space) and by the the celestial sphere where the equatorial plane
equation of state of the atmospheric mixture. intersects the ▶ ecliptic plane. When the Sun is at
Equations of state are also at the heart of our one of these points, daytime and nighttime have the
knowledge of planetary and stellar interiors. same duration everywhere on Earth. The term
ERA 745
equinox is also used for the date and time Overview

corresponding to those two positions of the Sun,
which mark the beginning of spring (vernal equi- ERA was mounted on the sun-pointing cold plate
nox) and the beginning of fall (autumnal equinox). of ESA’s European Retrievable Carrier
(▶ EURECA), which was launched to Low
Earth Orbit (LEO) on July 31, 1992, and
See Also remained there until August 7, 1993 (Innocenti
and Mesland 1995). ERA consisted of an expo-
▶ Coordinate Systems sure tray (Fig. 1) and a lid accommodating the
▶ Ecliptic following experiments:
E
▶ Right Ascension
• Exobiological unit that studied the action of
solar UV radiation and/or space vacuum on
the survival and genetic changes of inverte-
brates, ▶ microorganisms, viruses, and DNA
ERA
and the role of chemical and physical protec-
tion mechanisms (Horneck et al. 1995)
Gerda Horneck
• Space biochemistry that elucidated dehydra-
tion and photochemical reactions in cellular,
Aerospace Medicine, Radiation Biology, Köln,
subcellular, and molecular systems under
Germany
outer space conditions (Dose et al. 1995)
• Effects of solar UV on yeast cells
• Photoprocessing of grain mantle analogues
Definition
that studied photolytical processes during
chemical evolution of interstellar grains
The Exobiology Radiation Assembly (ERA) is
(Greenberg 2000)
a facility to expose various chemical and biolog-
ical objects to the conditions of outer space, such
Beneath the tray was another compartment
as space vacuum, solar extraterrestrial radiation,
accommodating the following experiment:
and cosmic rays.
ERA, Fig. 1 Tray of the

exposure facility ERA with
sample carriers and optical
filters during assembly
(Credit ESA, from Horneck
et al. 2010)
746 ERE
• Free Flyer ▶ Biostack that studied the biolog-

ical responses of different systems to the struc- Eros Asteroid
tured components of cosmic radiation, i.e.,
▶ HZE particles (cosmic rays consisting of Gerhard Hahn
energetic nuclei with atomic number 3 or Asteroids and Comets, DLR, Institute of
greater) and nuclear disintegration events Planetary Research, Berlin, Germany
(Reitz et al. 1995; Horneck 2007; Horneck
et al. 2010)
Synonyms
See Also
Asteroid (433) Eros
▶ Biostack
▶ EURECA
Definition
▶ Expose
▶ Exposure Facilities
(433) Eros is the second largest Amor asteroid, of
▶ HZE Particle
irregular shape measuring 34 11 11 km and
of spectral type S. Its rotation period is 5.27 h,
▶ Microorganism
exhibiting large brightness variations caused by
▶ Radiation Biology
the elongated shape. In 2000, NASA’s NEAR-
▶ Solar UV Radiation, Biological Effects
Shoemaker mission visited the asteroid, orbiting
▶ Space Environment
it for about 1 year, before descending to the
▶ Space Vacuum Effects
surface.
▶ Yeast
Dose K, Bieger-Dose A, Dillmann R, Gill M, Kerz O, (433) Eros was discovered on August 13, 1898,
Klein A, Meinert H, Nawroth T, Risi S, Stridde
by Gustav Witt in Berlin and almost simulta-
C (1995) ERA-experiment “Space biochemistry”.
Adv Space Res 16(8):119–129 neously by Auguste Charlois in Nice (see Scholl
Greenberg JM (2000) The secret of stardust. Sci Am and Schmadel (2002) for details). It was the first
283:46–51 near-Earth asteroid found.
Horneck G (2007) Space radiation biology. In:
Brinckmann E (ed) Biology in space and life on
Earth. Wiley-VCH, Weinheim, pp 243–273
Horneck G, Eschweiler U, Reitz G, Wehner J, Willimek R,
See Also
Strauch K (1995) Biological responses to space: results
of the experiment “Exobiological Unit” of ERA on
EURECA I. Adv Space Res 16(8):105–118 ▶ NASA
Horneck G, Klaus DM, Mancinelli RL (2010) Space ▶ Near-Earth Objects
microbiology. Microbiol Mol Biol Rev 74:121–156
Innocenti L, Mesland DAM (eds) (1995) EURECA scien-
tific results. Adv Space Res 16(8):1–140
Reitz G, Atwell W, Beaujean R, Kern JW (1995) Dosimet- References and Further Reading
ric results on EURECA. Adv Space Res
16(8):131–137 NEAR Shoemaker http://nssdc.gsfc.nasa.gov/planetary/
near.html. Last accessed 5 May 2014
Scholl H, Schmadel LD (2002) Discovery circumstances
of the first near-earth Asteroid (433) Eros. In: Dick
ERE WR, Hamel J (eds) Beiträge zur Astronomie-
geschichte, Band 5, Verlag Harry Deutsch, Thun and
▶ Extended Red Emission Frankfurt am Main, Germany, pp 210–220
Ester 747
to a larger mass. This is important for numerous

Error Rate processes: atmospheric molecules escaping from
a planet, a planet escaping from its host star,
Ester Lázaro and stars escaping from either mutual orbits or
Molecular Evolution Laboratory, Centro de stellar clusters. For a simple Keplerian potential
Astrobiologı́a (CSIC-INTA), Torrejón de Ardoz, (where the gravitational force is proportional
Madrid, Spain to the central mass and inversely proportional
to the distance squared), the escape speed is
given by
Definition E
rffiffiffiffiffiffiffiffiffiffiffi
M
In biology, the error rate is the average number of vesc ¼ 2G
▶ mutations per nucleotide and round of copy R
produced during genomic ▶ replication. For
example, an error rate of 1010 in a certain species where G is the gravitational constant, M is the
means that, in average, one in every 1010 nucleo- central mass, and R is the distance from the
tides will be misincorporated each time its genome central mass.
is replicated. The error rate varies by orders of
magnitude among species (from 103–105 in
RNA viruses to 109–1012 in cellular organ- See Also
isms). It also can present intraspecific variations,
depending on the nature and frequency of change ▶ Ejection, Hyperbolic
of the selective pressures. The optimal value of the
error rate results from the necessity to optimize
different conflicting features, such as the genera-
tion of genetic diversity, the preservation of
genetic information, the minimization of the num- Ester
ber of deleterious mutations, and the metabolic
costs of systems that increase replication fidelity. Henderson James (Jim) Cleaves II
See Also Institute of Technology, Meguro–ku, Tokyo,
Japan
▶ Mutation Institute for Advanced Study, Princeton, NJ,
▶ Natural Selection USA
▶ Replication (Genetics) Blue Marble Space Institute of Science,
Washington, DC, USA
Escape Velocity
Sean N. Raymond
Laboratoire d’Astrophysique de Bordeaux, Definition
In chemistry, esters are compounds derived by
Definition condensation of an oxoacid (one containing an
oxo group, X=O, e.g., phosphoric acid (O=P
Escape velocity is the critical velocity above (OH)3) or a carboxylic acid (RCOOH)) with an
which a body is no longer gravitationally bound alcohol. Many biological lipids are the fatty acid
748 Eta-Earth
esters of glycerol derivatives. Phosphoesters

form the backbone of DNA molecules. Polyes- Eta-Earth
ters such as poly-hydroxy butyrate are impor-
tant energy storage molecules in some Nader Haghighipour
microbes. Cyclic esters formed from carboxylic Institute for Astronomy, University of Hawaii-
acid and alcohol functional groups are called Manoa, Honolulu, Hawaii, HI, USA
lactones.
Definition
Overview The term eta-Earth (also written as ZE) is defined

as the mean number per star of rocky planets with
Esters contain a carbonyl functionality, but between 1 and 1.5–2 Earth-radii that reside in the
unlike amides, esters are structurally flexible optimistic habitable zone (HZ) of their host star.
because there is a low energetic barrier to rota- Eta-Earth enters one formulation of the ▶ Drake
tion about the X(=O)–O–C bond. Esters thus equation, which endeavors to estimate the occur-
tend to be more volatile (i.e., have a lower boil- rence of intelligent life in the Galaxy; at the
ing point, particularly with low molecular present time, it is usually calculated separately
weight esters) than the corresponding amides. for each stellar spectral type. Thus, eta-Earth
Their carbonyl group can serve as a hydrogen- represents the occurrence rate of rocky planets
bond acceptor, and this ability to engage in in the optimistic HZ of different stars.
hydrogen bonding renders them somewhat The References present some values for eta
water soluble. The inability of the linking ester Earth based on different statistical analyses of
oxygen atom to engage in hydrogen bonding the data from the Kepler space telescope.
with another ester group, unlike the protonated
nitrogen atom in an amide or a protonated car-
boxylic acid, also results in their greater struc- See Also
tural flexibility and volatility, but renders them
unable to form the hydrogen-bonded structural ▶ Exoplanets, Discovery
motifs such as a-helices and b-sheets that ▶ Habitable Zone
polyamides can.
Esters can react at one of two locations
of their molecular structure. The carbonyl
group is weakly electrophilic and is attacked Dressing CD, Charbonneau D (2013) The occurrence rate
by strong nucleophiles (amines, alkoxides, of small planets around small stars. Astrophys
etc.). The C–H bonds of the carbon atoms adja- J 767(1). Article id. 95
cent to the carbonyl group are weakly acidic but Gaidos E 2013 Candidate planets in the habitable zones of
Kepler stars. Astrophys J 770(2). Article id. 90
can be deprotonated by strong bases and can Kopparapu RK 2013 A revised estimate of the occurrence
thus serve as nucleophiles. Esters undergo rate of terrestrial planets in the habitable zones around
both acid and base catalyzed hydrolysis. Under Kepler M-dwarfs. Astrophys J Lett 767(1). Article
basic conditions, hydroxide acts as a nucleoid. L8
phile, while an alkoxide is the leaving group.
This reaction is known as saponification. Esters
may also be cleaved by amines such as ammonia
and primary or secondary amines to give Ethanal
amides, in a process known an aminolysis or
amidation. ▶ Acetaldehyde
Ethanol 749
Loomis RA, Zaleski DP, Steber AL, Neill JL, Muckle MT,
1,2-Ethanediol Harris BJ, Hollis JM, Jewell PR, Lattanzi V, Lovas FJ,
Martinez O, McCarthy MC, Remijan AJ, Pate BH,
Corby JF JF (2013) The detection of interstellar
▶ Ethylene Glycol ethanimine (CH3CHNH) from observations taken dur-
ing the GBT PRIMOS survey. Astrophys J 765:
L9–L16
Ethanimine
William M. Irvine E
University of Massachusetts, Amherst, MA, USA Ethanoic Acid
▶ Acetic Acid
Synonyms
CH3CHNH
Ethanol
Definition
Kensei Kobayashi
Ethanimine is an 8-atom, nitrogen-containing Yokohama National University, Tokiwadai,
organic molecule that is the next heavier homo- Hodogayaku, Yokohama, Japan
logue of methanimine (CH2NH). In laboratory
experiments involving the irradiation of interstel-
lar ice analogs, it has been suggested that Synonyms
ethanimine is a precursor to the formation of the
amino acid alanine. Ethyl alcohol
History Definition
Methanimine was detected in the interstellar Ethanol is an organic compound belonging to the
medium in 1973 (Godfrey et al. 1973), and group of ▶ alcohols whose structural formula is
ethanimine has more recently been found in the C2H5OH. It is a constituent of alcoholic bever-
Milky Way’s galactic center molecular cloud ages. Ethanol is a colorless, volatile, and flam-
Sagittarius B2 North (Loomis et al. 2013). mable liquid. Its melting and boiling points are
114.3 C and 78.4 C, respectively. It is made
from sugar by fermentation, and it is metabo-
See Also lized in vivo to carbon dioxide via acetaldehyde
and acetic acid. Ethanol has rotational isomers,
▶ Molecules in Space and they are referred to as trans-ethanol and
gauche-ethanol (Fig. 1). In 1975, trans-ethanol
was identified as an interstellar molecule,
References and Further Reading while gauche-ethanol was detected in 1996. Eth-
Godfrey PD, Brown RD, Robinson BJ, Sinclair MW
anol condenses with carboxylic acids to form
(1973) Discovery of interstellar methanimine esters, and it condenses with itself to form
(formaldimine). Astrophys Lett 13:119–121 diethyl ether.
750 Ether
Ethanol,
Fig. 1 Rotational isomers
of ethanol
See Also (five-membered rings) and epoxides (three-

membered rings).
▶ Alcohol Ethers cannot form intermolecular hydrogen
▶ Methanol bonds between each other, resulting in relatively
▶ Molecular Cloud low boiling points compared to their parent alco-
hols. The difference in the boiling points of ethers
and their parent alcohols becomes less significant
as the carbon chains become longer, as van der
Waals interactions between the extended carbon
Ether
chains come to dominate intermolecular
interaction.
Ethers in general are of low chemical reactiv-
ity, and indeed in many extremophilic organisms,
the cell membranes contain glycerol ethers rather
Japan
than glycerol esters.
USA
Washington, DC, USA
Center for Chemical Evolution, Georgia Institute Ethyl Alcohol
▶ Ethanol
Definition
Ethers are a class of organic compound

containing an ether group (R-O-R) – an oxygen Ethyl Cyanide
atom connecting two alkyl or aryl groups. A typ-
ical example is diethyl ether, commonly referred William M. Irvine
to simply as “ether” (CH3-CH2-O-CH2-CH3). University of Massachusetts, Amherst, MA, USA
Polyethers are compounds with more than one
ether group. Low to medium range molecular
weight polyethers with a hydroxyl end group Synonyms
are termed glycols. The crown ethers are exam-
ples of low-molecular weight cyclic polyethers. CH3CH2CN; Cyanoethane; Propanenitrile
Other common cyclic ethers include the furans (IUPAC name); Propionitrile
Ethyl Formate 751
Definition
Ethyl Formate
Under standard laboratory conditions, ethyl cya-
nide is a clear liquid with a sweet odor. It is William M. Irvine
commonly found in the gas phase in interstellar University of Massachusetts, Amherst, MA, USA
molecular clouds, specifically in “hot cores” (the
regions where massive stars are forming), and has
more recently been located in regions of Synonyms
low-mass star formation (hot corinos). Ethyl cya-
nide has a rich rotational spectrum that is C2H5OCHO; Ethyl methanoate
E
observed by radio astronomers in the millimeter
wavelength region. In addition to transitions in
the ground vibrational state, rotational lines in Definition
low-lying vibrational states have been detected.
All three 13-carbon isotopic species have been Ethyl formate is among the largest organic mol-
detected in hot cores. ecules unambiguously detected by radio astrono-
mers in the interstellar medium. It is also called
ethyl methanoate. Ethyl formate is partly respon-
History sible for the flavor of raspberries, and it has the
characteristic odor of rum.
The first astronomical detection was made by
Johnson et al. (1977); subsequently it has been
found in most spectral surveys of hot cores (e.g.,
History
Nummelin et al. 2000; Daly et al. 2013).
The trans conformer of ethyl formate was
detected in the Galactic Center molecular cloud
See Also Sagittarius B2(N) by Belloche et al. (2009), who
suggest that it may be produced by surface reac-
▶ Hot Core tions on interstellar dust grains, perhaps involv-
▶ Hot Corino ing precursors such as methyl formate and
▶ Isotope ethanol (for the definition of conformers, see
▶ Molecular Cloud ▶ isomer). More recently, the gauche conformer
▶ Molecules in Space has also been detected, in a line survey of the
Orion molecular cloud (Tercero et al. 2013).
References and Further Reading Ethyl formate is the second formate, after
▶ methyl formate, to be detected in the ▶ inter-
Daly AM, Bermúdez C, López A, Tercero B, Pearson JC, stellar medium. Its ▶ isomer ▶ methyl acetate
Marcelino N, Alonso JL, Cernicharo J (2013) Labora- (CH3COOCH3) has also been recently detected
tory characterization and astrophysical detection of
in a molecular cloud in the Milky Way.
vibrationally excited states of ethyl cyanide. Astrophys
J 768:81
Johnson DR, Lovas FJ, Gottlieb CA, Gottlieb EW,
Litvak MM, Thaddeus P, Guélin M (1977) Detection
of interstellar ethyl cyanide. Astrophys J 218: See Also
L370–L376
Nummelin A, Bergman P, Hjalmarson Å, Friberg P, Irvine ▶ Interstellar Medium
WM, Millar TJ, Ohishi M, Saito S (2000) A three-
position spectral line survey of sagittarius B2 between
▶ Methyl Acetate
218 and 263 GHZ. II. Data analysis. Astrophys J Suppl ▶ Methyl Formate
128:213–243 ▶ Molecules in Space
752 Ethyl Methanoate
References and Further Reading center. This contrasts with the relative strength of
the lines in comet C/1995 O1 (Hale-Bopp), where
Belloche A, Garrod RT, M€ uller HSP, Menten KM, this molecule was identified in archive spectra as
Comito C, Schilke P (2009) Increased complexity in
soon as the line frequencies were available
interstellar chemistry: detection and chemical model-
ing of ethyl formate and n-propyl cyanide in Sagittar- (Crovisier et al. 2004). Several conformers of
ius B2(N). Astron Astrophys 499:215–232 ethylene glycol exist (Hollis et al. 2002), and
Tercero B, Kleiner I, Cernicharo J, Nguyen HVL, the astronomical detections prior to 2012 referred
López A, Muñoz Caro GM (2013) Discovery of
to the lowest energy conformer. A second con-
methyl acetate and gauche ethyl formate in Orion.
Astrophys J 770:L13–L19 former has recently been reported (2013) in the
interstellar medium in as yet unpublished
research.
Ethyl Methanoate
See Also
▶ Ethyl Formate
▶ Comet
▶ Ethylene Oxide
Ethylene Glycol
Didier Despois
Laboratoire d’Astrophysique de Bordeaux, Crovisier J, Bockelée-Morvan D, Biver N, Colom P,
Despois D, Lis DC (2004) Ethylene glycol in comet
CNRS-Universite de Bordeaux, France
C/1995 O1 (Hale-Bopp). Astron Astrophys 418:
L35–L38
Hollis JM, Lovas FJ, Jewell PR, Coudert LH (2002) Inter-
Synonyms stellar antifreeze: ethylene glycol. Astrophys J Lett
571:L59–L62
1,2-Ethanediol; Glycol; HOCH2CH2OH
Definition
Ethylene Oxide
Ethylene glycol [IUPAC name 1,2-ethanediol] is
William M. Irvine
the simplest dialcohol (or diol). A (toxic) liquid
under standard pressure from 197 C down to
12 C, its mixture (70–30 %) with water
remains liquid down to 51 C, which explains
its use as antifreeze. It is currently produced by Synonyms
reaction of ▶ ethylene oxide (oxirane, C2H4O)
with water. Ethylene glycol has been found in C2H4O; Oxirane
the interstellar medium toward the galactic center
and in comet C/1995 O1 (Hale-Bopp).
Definition
History
Detection of ethylene glycol in interstellar matter Under laboratory conditions, ethylene oxide is
was rather difficult; it was performed for the first a colorless, flammable gas. Ethylene oxide
time by Hollis et al. (2002) toward the galactic (IUPAC name oxirane) is one of only a handful
Ethynyl Radical 753
of cyclic molecules that have been found in the

interstellar medium, all of which except benzene Ethyl-glycine
involve three-membered rings. It was detected by
radio astronomers by observing pure rotational ▶ Aminobutyric Acid
transitions at millimeter wavelengths in the
Galactic Center ▶ hot core SgrB2(N). Ethylene
oxide is isomeric with ▶ acetaldehyde
(CH3CHO) and vinyl alcohol (CH2CHOH), both
of which are also observed in the interstellar
Ethyne, HCCH
medium. NASA has used ethylene oxide for ster-
▶ Acetylene E
ilization of spacecraft as part of its ▶ planetary
protection program.
History Ethynyl Radical
Ethylene oxide was isolated in 1859 by the William M. Irvine

French chemist C.A. Wurtz. It subsequently University of Massachusetts, Amherst, MA, USA
became commercially important as a precursor
to ethylene glycol, and it was used during World
War I in the production of mustard gas. The
Synonyms
astronomical detection was carried out after
a failed search for its derivative, oxiranecar-
C2H
bonitrile (C3H3NO), which had been suggested
as a possible precursor to glycolaldehyde phos-
phate and hence to sugar phosphates (Dickens
et al. 1996). Definition
This triatomic ▶ radical plays an important role

See Also in the gas phase chemistry of interstellar
▶ molecular clouds, since it provides a link in
▶ Acetaldehyde the chemistry of acetylene and higher-order
▶ Hot Core polyacetylenes, the presence of the latter being
▶ Isomer deduced from observations of the corresponding
▶ Molecular Cloud nitriles (▶ cyanopolyyne). Both carbon-13 and
▶ Molecules in Space deuterated isotopic variants of C2H are observed
▶ Planetary Protection in ▶ molecular clouds.
History
Dickens JE, Irvine WM, Ohishi M, Arrhenius G, Pitsch S,

The fundamental pure rotational transition
Bauder A, M€uller F, Eschenmoser A (1996) A search (N = 1 0) of C2H at a frequency of 87 GHz
for interstellar oxiranecarbonitrile (C3H3NO). Orig was observed astronomically before
Life Evol Biosph 26:97–110 corresponding laboratory measurements were
Dickens JE, Irvine WM, Ohishi M, Ikeda M, Ishikawa S,
available, with the identification resting on the
Nummelin A, Hjalmarson Å (1997) Detection of inter-
stellar ethylene oxide (c-C2H4O). Astrophys pattern of the four detected hyperfine components
J 489:753–757 (Tucker et al. 1974).
754 Eucarya
See Also features including but not limited to nuclear pores,

endoplasmic reticulum, 9 + 2 flagellar apparatus,
▶ Cyanopolyyne mitotic spindle formation, acidified vacuoles,
▶ Deuterium Golgi apparatus, multiple linear chromosomes,
▶ Molecular Cloud eukaryotic telomeres, and cellular processes that
▶ Molecules in Space typically include mitosis, meiotic sex, endocytosis,
▶ Radical and mitochondrial respiration. They are also dif-
ferentiated from the other two domains by the
presence of specific genes and proteins (e.g., tubu-
lins, actin, dyneins, centrin, myosin, calmodulin,
Tucker KD, Kutner ML, Thaddeus P (1974) The ethynyl and ubiquitin).
radical C2H – a new interstellar molecule. Astrophys
J 193:L115–L119
Overview
Members of the domain Eukarya include both

Eucarya unicellular and multicellular representatives from
the 1-mm ocean-dwelling “picoplanktonic” alga
▶ Eukarya Ostreococcus to the blue whale (34 m) – a differ-
ence in size of over 7 orders of magnitude. Ani-
mals, plants, and ▶ fungi all have microbial
eukaryotic (protistan) sister groups. Traditional
Eukarya taxonomic classifications of the Eukarya are mor-
phology based and have described 1.25 million
Linda Amaral-Zettler animal species (mostly insects), 297,326 plant spe-
Marine Biological Laboratory, Josephine Bay cies and 75,000 fungal species, and 200,000
Paul Center for Comparative Molecular Biology protistan species (http://www.eol.org/). Single-
and Evolution, Woods Hole, MA, USA cell and some multicellular protists contribute to
the greatest diversity of life on Earth in the domain
Eukarya. Beginning in the 1960s, a large portion of
Keywords this diversity was unveiled with the assistance of
technologies including high-powered electron
Classification; Domain; Tree of life microscopy. Unlike bacteria and archaea, many
microbial eukaryotes have ultrastructural features
that can help to differentiate the major groups.
Synonyms These ultrastructural features are summarized in
below (Patterson 1999).
Eucarya; Eukaryote Ultrastructural features that differentiate
major groups of eukaryotes are:
Definition • Shape of mitochondrial cristae (i.e., tubular,

elongated, branching, flat, or discoidal)
Eukarya is one of the three domains of life distin- • Presence or absence of hairs, scales, or other
guished from ▶ Bacteria and ▶ Archaea at the extensions on the flagella
morphological and molecular levels. All members • Organization of the axoneme into the nine-
of the Eukarya have a nucleus and are further triplet structure of the basal bodies
distinguished from Bacteria and Archaea by a • Basal body length and orientation association
complex cellular organization with ultrastructural with other organelles
Eukarya 755
• Microtubule and rootlet structures arising best. Another holdfast has been the prokaryote-
from the basal bodies eukaryote dichotomy. Pace (2006) pointed out
• Source and deployment of microtubular or the need to recognize the value of the three-
other cytoskeletal arrays within the cell domain system in doing away with the artificial
• Presence and nature of microtubule- lumping of bacteria and archaea via the term
organizing centers “prokaryote.” While ribosomal RNA approaches
• Number, nature, and heterogeneity of nuclei, failed to unveil higher-order relationships
structures within the nuclear envelope, between the three domains, amino acid-based
intranuclear and paranuclear inclusions analyses using vacuolar H+-ATPases showed
• Behavior of the nuclear envelope during that Archaea are more closely related to Eukarya
E
mitosis than they are to Bacteria (Gogarten et al. 1989),
• Behavior of the mitotic spindle further obfuscating the prokaryote-eukaryote
• For chloroplast-containing cells: number of division.
bounding membranes, thylakoids per lamella, While important in establishing lower-level
and the presence and nature of contained (e.g., taxonomic relationships, single-gene approaches
stigma) or associated (e.g., nucleomorph) such as rRNA, tubulin, and others often returned
organelles ambiguous relationships between major groups
• Identity and nature of other membrane-bound of eukaryotes. The adoption of multigene phylo-
organelles in the cell genetic approaches to inferring relationships
between eukaryotes, followed by the use of geno-
Based on these ultrastructural features, mic and expressed sequence tag (EST) data, ush-
Patterson (1999) defined 71 groups of protists ered in the “phylogenomic” era in eukaryotic
but identified an additional 200 with no clear phylogeny research. These approaches allowed
ultrastructural identity. While ultrastructure has for the testing of relationships between major
its virtues in the eukaryotic domain, molecular lineages of eukaryotes and lead to the adoption
approaches quickly superseded morphological of the “supergroup” concept of eukaryotic phy-
ones in accelerating our ability to delineate spe- logeny that encompasses a varying number of
cies and higher-order taxa of eukaryotes. After higher taxonomic groupings than phyla
Woese and Fox’s (1977) seminal work on apply- depending on the extent of the supergroup.
ing ribosomal RNA-based strategy for inferring These include the Opisthokonta, Amoebozoa,
the relationships between the major branches of Archaeplastida (Plantae), Rhizaria,
the tree of life, this approach quickly became the Chromalveolata (chromalveolates, chromists
gold standard in molecular systematics studies in plus alveolates), and the Excavata (Baldauf
the 1980s and 1990s across all domains of life. In 2003; Keeling et al. 2005; Simpson and Roger
1990, Woese et al. (1990) formally erected the 2004).
three domains including the Bacteria, Archaea, More detailed information on these super-
and Eukarya. Many studies employing rRNA- groups can be found in the references listed
based approaches ensued and made important above, but a brief description follows. The
contributions to the placement of eukaryotes Opisthokonta encompass the animal and fungal
into major groups (Sogin 1991). kingdoms, as well as several groups of protists
Despite insights gained in applying molecular related to both of these lineages, choanofla-
metrics to redefining the organization of life, gellates and Mesomycetozoea (a group of para-
modern taxonomic revisions have been slow to sitic protists, many in fishes), branch with the
reach textbooks. The animal-plant dichotomy animals, and nucleariids and microsporidia
was replaced with the Whittaker five kingdoms (a group of obligate endoparasites) branch with
recognizing Monera, Animalia, Plantae, Fungi, the Fungi. Their name reveals the nature of the
and Protista, but the transition to the three- synapomorphy or shared derived trait that its
domain classification has been a gradual one at members share – a single posterior flagellum.
756 Eukarya
This higher-order relationship between two The Chromalveolata supergroup itself con-
kingdom-level eukaryotic groups has been tains higher-order groupings of eukaryotes
known for some time and was first proposed on including the alveolates (i.e., ciliates,
the basis of morphological characters (Cavalier- apicomplexans, dinoflagellates) and chromists
Smith 1987), confirmed via single-gene rRNA- (Stramenopiles – brown algae, diatoms,
based molecular analyses (Wainright et al. 1993) pelagophytes, labyrinthulids, oomycetes,
and more recently supported by various bicosoecids, opalindids), haptophytes, and
phylogenomic investigations (Burki et al. 2007; cryptomonads. Members of this supergroup are
Hampl et al. 2009; Parfrey et al. 2006). ecologically diverse ranging from free-living to
The Amoebozoa form a second supergroup obligately parasitic and heterotrophic, autotro-
comprising lobose amoebae, slime molds, as phic, and mixotrophic. The Chromalveolates
well as some amitochondriate amoebae including remains among the most controversial of super-
pelobionts and entamoebae (sometimes referred groups and may require more taxon sampling to
to as Archamoebae). Like the terms “alga” and justify (Hampl et al. 2009).
“worm,” “amoeba” joins the ranks of ecological The Rhizaria consists of unicellular and
concepts as opposed to phylogenetic ones colony-forming (therefore multicellular) pro-
because the term does not refer to a coherent tists that include the geologically significant
group of organisms that share common ancestry. Radiolaria and Foraminifera that are well pre-
Amoeboid organisms, organisms that possess served in the fossil record, as well as a hodge-
pseudopodia or “false feet” that enable the cell podge of other amoeboid and flagellated
to engulf food via phagocytosis, are sprinkled forms collectively referred to as the Cercozoa.
over the tree of life and occur in the Opisthokonta Members of the Radiolaria have been detected in
(i.e., nucleariids), Amoebozoa, Rhizaria (i.e., molecular studies targeting picoplankton-sized
Foraminifera, Polycystines, Phaeodaria, (<2 mm) cells, as well as colonies that reach
Acatharea), and Excavata (i.e., Heterolobosea). several meters in length. In addition to
Amoebozoa include free-living, facultative, and exhibiting extreme morphological variability,
obligate parasites including the genus Amoeba, like the Chromalveolates, the supergroup also
Acanthamoeba, and Entamoeba. As with the exhibits varied ecological roles including het-
Opisthokonta, Amoebozoa have multicellular erotrophic, autotrophic, and parasitic lifestyles,
representatives including members of the slime and many members of the Radiolaria and Fora-
molds such as Dictyostelium. minifera in particular possess symbiotic algae.
The Plantae supergroup, also called the Rhizaria is defined solely on the basis of molec-
Archaeplastida, includes unicellular and ular phylogenies and lacks any synapomorphic
multicellular members such as the land plants, characters.
charophytes, chlorophytes, red ▶ algae, and The final supergroup is the Excavata
glaucophytes. All the members of this group are (Patterson 1999) that include the Heterolobosea,
photosynthetic or contain relict plastids resulting euglenids, diplonemids, kinetoplastids, jacobids,
from a primary endosymbiosis with a cyanobac- Carpediemonas, diplomonads, retortamonads,
terium. Surprisingly, a recent phylogenomic trichomonads, hypermastigotes, Trimatix, and
assessment of this supergroup along with other the oxymonads. Phylogenomic analyses have
supergroups including 143 proteins and 48 taxa only recently supported the monophyly of this
failed to support the monophyly of the supergroup (Hampl et al. 2009), which, unlike
Archaeplastida (Hampl et al. 2009). Beyond sys- other supergroups, was primarily conceived on
tematic errors associated with the analysis, one the basis of an ultrastructural feature of a distinc-
possible explanation is related to the symptom- tive ventral feeding groove occurring in the flag-
atic problem of the host nuclear genome being ellated members of the group and further
a potential mosaic of genes derived from multiple expanded via molecular studies. The Excavata
nuclei (Lane and Archibald 2008). contains many human disease-causing
Eukaryote 757
representatives including Giardia (diplomonad), Baldauf SL (2003) The deep roots of eukaryotes. Science
Naegleria (heterolobosean), and Trypanosoma 300:1703–1706
Burki F, Shalchian-Tabrizi K, Minge M, Skjaeveland A,
(kinetoplastid), as well as many free-living non- Nikolaev SI, Jakobsen KS et al (2007) Phylogenomics
pathogenic members. Among them is Euglena reshuffles the eukaryotic supergroups. PLoS One 2:
that includes both heterotrophic and autotrophic e790
representatives. As with other supergroups, the Cavalier-Smith T (1987) The origin of fungi and
pseudofungi. In: Rayner ADM (ed) Evolutionary biol-
Excavata also contains members that are ogy of the fungi. Cambridge University Press, Cam-
multicellular, namely, heterolobosean acrasids bridge, pp 339–353
that form a sporangia much like the slime molds Gogarten JP, Kibak H, Dittrich P, Taiz L, Bowman EJ,
of the Amoebozoa – but phylogenetically dis- Bowman BJ et al (1989) Evolution of the vacuolar H+
-ATPase: implications for the origin of eukaryotes. E
tinct. Also like slime molds, they serve as major Proc Natl Acad Sci U S A 86:6661–6665
decomposers in the environment. Hampl V, Hug L, Leigh JW, Dacks JB, Lang BF, Simpson
AGB et al (2009) Phylogenomic analyses support the
monophyly of Excavata and resolve relationships
among eukaryotic “supergroups”. Proc Natl Acad Sci
Future Directions U S A 106:3859–3864
Keeling PJ, Burger G, Durnford DG, Lang BF, Lee RW,
With increasing numbers of rRNA-based envi- Pearlman RE et al (2005) The tree of eukaryotes.
ronmental clone surveys unveiling potentially Trends Ecol Evol 20:670–676
Lane CE, Archibald JM (2008) The eukaryotic tree of life:
novel eukaryotic groups, there is still the chance endosymbiosis takes its TOL. Trends Ecol
that there remain unexplored supergroups of the Evol 23:268–275
domain Eukarya (Massana and Pedros-Alio Massana R, Pedros-Alio C (2008) Unveiling new micro-
2008). Among the possibilities are sequences bial eukaryotes in the surface ocean. Curr Opin
related to Apusomonads and Picobiliphyta, two Pace NR (2006) Time for a change. Nature 441:289–289
protist groups that do not have clear affinities to Parfrey LW, Barbero E, Lasser E, Dunthorn M,
existing supergroups. New technologies afforded Bhattacharya D, Patterson DJ et al (2006) Evaluating
by next-generation pyrosequencing (Amaral- support for the current classification of eukaryotic
diversity. PLoS Genet 2:e220
Zettler et al. 2009) and single-cell ▶ genomics Patterson DJ (1999) The diversity of eukaryotes. Am Nat
approaches hold promise for completing our 154:S96–S124
search for novelty among the Eukarya. Roger AJ (1999) Reconstructing early events in eukary-
otic evolution. Am Nat 154:S146–S163
Simpson AGB, Roger AJ (2004) The real “kingdoms” of
eukaryotes. Curr Biol 14:R693–R696
Sogin ML (1991) Early evolution and the origin of eukary-
See Also
otes. Curr Opin Genet Dev 1:457–463
Wainright PO, Hinkle G, Sogin ML, Stickel SK
▶ Algae (1993) Monophyletic origins of the metazoa: an evo-
▶ Fungi lutionary link with fungi. Science 260:340–342
Woese CR, Fox GE (1977) Phylogenetic structure of the
▶ Genomics
prokaryotic domain: the primary kingdoms. Proc Natl
▶ Multicellular Organisms Acad Sci U S A 74:5088–5090
▶ Ribosome Woese CR, Kandler O, Wheelis ML (1990) Towards a
▶ Sequence Analysis natural system of organisms: proposal for the domains
archaea, bacteria, and eucarya. Proc Natl Acad Sci
▶ Yeast
U S A 87:4576–4579
Amaral-Zettler LA, McCliment EA, Ducklow HW, Huse

SM (2009) A method for studying protistan diversity
using massively parallel sequencing of V9 hypervari-
Eukaryote
able regions of small-subunit ribosomal RNA genes.
PLoS One 4:e6372 ▶ Eukarya
758 Eukaryotes, Appearance and Early Evolution of
a nucleated proto-eukaryote or even a viral origin

Eukaryotes, Appearance and Early for the ▶ nucleus (review in López-Garcı́a
Evolution of et al. 2006; Roger and Hug 2006).
The early eukaryotic fossil record includes car-
Emmanuelle J. Javaux bonaceous compressions (the organisms are pre-
Palaeobiogeology-Palaeobotany- served as a thin film of carbon); ▶ acritarchs
Palaeopalynology, Geology Department, (organic-walled vesicles with unknown biological
Université de Liège, Liège, Belgium affinities); multicellular organic-walled organ-
isms; vase-shaped microfossils; molds and casts
in sandstone, carbonate, phosphate, or shale; min-
Keywords eralized skeletons, walls, or scales; and phyloge-
netically informative molecules (molecular
Acritarchs; Eucarya; Evolution; Precambrian; biomarkers) (review in Javaux 2011). The origin
Protists of the domain Eucarya is not constrained and
might be an Archean event. However, the Archean
eukaryotic record is debated and limited to bio-
Definition markers, possibly indicating the presence of cells
able to synthesize eukaryotic sterols in their flex-
Eukaryotes are the organisms belonging to the ible membranes. Recently, this molecular record
phylogenetic domain Eucarya. Eukaryotic cells has been reassessed as contaminants (Rasmussen
are less diverse than ▶ prokaryotes (phylogenetic et al. 2008; French et al. 2013). The earliest
domains ▶ Archaea and ▶ Bacteria) metaboli- eukaryotic fossils are possibly 1.9 Ga macroscopic
cally, but have a complex cellular architecture coiled filaments (Grypania) preserved as carbona-
comprising a ▶ nucleus, a cytoskeleton ceous compressions and unambiguously 1.8 Ga
(a proteinaceous network structuring the cyto- acritarchs with walls ornamented by concentric
plasm to facilitate intracellular traffic, endo- and striations. In the Mesoproterozoic (around
exocytosis, and amoeboid locomotion), an 1.5 Ga), the eukaryotic microfossil record shows
endomembrane system (a system of internal higher diversity and more obvious ecological het-
membranes subdivided into several organelles erogeneity. The first process-bearing acritarch
and used for synthesis, processing, packaging, (Tappania plana, Fig. 1) appears and is interpreted
and transport of macromolecules such as lipids as eukaryotic based on the combined characters of
and ▶ proteins), and organelles such as mito- its complex ornamentation, large size, and decay-
chondria (or derived organelles) and ▶ chloro- resistant wall (Javaux et al. 2001). It might be as
plasts in photosynthetic eukaryotes. old as 1.7 Ga, based on recent revision of the
Ruyang group, in China, which includes also
other process-bearing and ornamented acritarchs
Overview (Pang et al. 2013). This and other associated early
microfossils display characters of a eukaryotic
Unicellular eukaryotes are called protists. They degree of organization and are interpreted as
are very diverse and occur in all the supergroups eukaryotes with a sophisticated cytoskeleton and
of the eukaryotic tree (Baldauf 2008). a nucleus (review in Knoll et al. 2006). The earli-
Multicellular eukaryotes include fungi, animals, est eukaryote that can be related to an extant clade
and plants. At present, the origin of the domain is a multicellular red alga (Bangiomorpha) dated
Eucarya is still unresolved. Several authors have at about 1.2 Ga (Butterfield 2000). From the late
proposed a range of hypotheses involving an Mesoproterozoic through the middle
association between ▶ prokaryotes (one Neoproterozoic (1.2–0.63 Ga), (stem and/or
archaeon and one bacterium) as the origin of crown) members of all extant supergroups but
eukaryotic cells. Others suggest the existence of one are recorded in the fossil record (example of
Eukaryotes, Appearance and Early Evolution of 759
Eukaryotes, Appearance
and Early Evolution of,
Fig. 1 Examples of early
eukaryotes. Left: Tappania
plana, a spiny acritarch
(about 160 mm in diameter)
interpreted as an early
eukaryote from the 1.5 Ga
Roper Group, Australia
(Photo E Javaux). Right:
Bangiomorpha, a
multicellular red algae,
from the 1.2 Ga Hunting E
Formation, arctic Canada
(Photograph courtesy of
N Butterfield)
molecular clock study in Douzery et al. 2004; ▶ Prokaryote

review of early eukaryotic diversification in ▶ Proterozoic Eon
Javaux 2011). During that time period, simulta- ▶ Tree of Life
neously to these important biological innovations
and changing ecology, changes in ocean chemistry
and the formation and breakup of the superconti- References and Further Reading
nent ▶ Rodinia occurred, all events probably
Baldauf S (2008) An overview of the phylogeny and
playing a role in eukaryote diversification diversity of eukaryotes. J Syst Evol 46:263–273
(review in Javaux 2011). Major biological inno- Butterfield NJ (2000) Bangiomorpha pubescens n. gen.,
vations such as multicellularity (Butterfield 2009), n. sp.: implications for the evolution of sex, multicel-
sex, biomineralization, heterotrophy (Porter lularity and the Mesoproterozoic/Neoproterozoic radi-
ation of eukaryotes. Paleobiology 26:386–404
2006), and eukaryotic photosynthesis appear, Butterfield NJ (2009) Modes of pre-Ediacaran multicel-
leading to ecological tiering and complex food lularity. Precambrian Res 173:201–211
webs and interactions. Complex multicellularity Cavalier-Smith T (2009) Megaphylogeny, cell body plans,
(tissue-grade then organ-grade organization) and adaptive zones: causes and timing of eukaryote basal
radiations. J Eukaryot Microbiol 5:26–33
animal biomineralization and predation evolved Douzery EJP, Snell EA, Bapteste E, Delsuc F, Philippe
during the ▶ Ediacaran (635–542 Ma). H (2004) The timing of eukaryotic evolution: does a
relaxed molecular clock reconcile proteins and fossils?
Proc Natl Acad Sci U S A 101:15386–15391
French K, Hallan C, Hope J, Buick R, Brocks J, Summons
See Also R (2013) Archean hydrocarbon biomarkers: Archean
or not? Mineral Mag 77(5):110
▶ Acritarch Javaux EJ (2011) Evolution of early eukaryotes in Pre-
▶ Archaea cambrian oceans. In: Gargaud M, Lopez-Garcia P,
Martin H, Montmerle Th (eds) Origin of life: an astro-
▶ Bacteria biology perspective. Cambridge University Press,
▶ Chloroplast pp 414–449
▶ Cytoplasm Javaux EJ, Knoll AH, Walter MR (2001) Morphological
▶ Mitochondrion and ecological complexity in early eukaryotic ecosys-
tems. Nature 412:66–69
▶ Nucleus Knoll AH (2011) The multiple origins of complex
▶ Phylogeny multicellularity. Annu Rev Earth Planet Sci
▶ Precambrian 39:217–239
760 EURECA
Knoll AH, Javaux EJ, Hewitt D, Cohen P (2006) Eukary- was ERA (Exobiology and Radiation Assembly)
otic organisms in proterozoic oceans. Philos Trans in which bacteria and organic materials were
R Soc B 361:1023–1038
López-Garcı́a P, Moreira D, Douzery E, Forterre P, van exposed to study their survival and evolution in
Zuilen M, Claeys P, Prieur D (2006) Ancient fossil space.
record and early evolution (ca. 3.8–0.5 Ga). Earth It was a precursor to the ▶ BIOPAN and
Moon Planet 98:247–290 ▶ EXPOSE facilities.
Pang K, Tanq Q, Schiffbauer JD, Yao J, Yuan, Wan B,
Chen L, Ou Z, Xiao S (2013) The nature and origin of
nucleus-like intracellular inclusions in Paleopro-
terozoic eukaryote microfossils. Geobiology
11:499–510 See Also
Porter SM (2006) The proterozoic fossil record of hetero-
trophic eukaryotes. In: Xiao S, Kaufman AJ (eds) ▶ BIOPAN
Neoproterozoic geobiology and paleobiology. ▶ EXPOSE
Springer, Dordrecht, pp 1–21
Rasmussen B, Fletcher IR, Brocks JJ, Kilburn MR ▶ Exposure Facilities
(2008) Reassessing the first appearance of eukaryotes
and cyanobacteria. Nature 4555:11101–11105
Roger AJ, Hug LA (2006) The origin and diversification
of eukaryotes: problems with molecular phylogenetics
and molecular clock estimation. Philos Trans R Soc Europa
B 361:1039–1054
Therese Encrenaz
EURECA
Hervé Cottin Keywords

Laboratoire Interuniversitaire des Systèmes
Atmosphériques, Université Paris Est-Créteil, Galilean satellites
Créteil, France
Definition
Definition
Europa, discovered by ▶ Galileo Galilei in 1610,
EURECA (EUropean REtrievable CArrier) was is the smallest of the four Galilean satellites
a 4.5-t satellite designed to carry spaceborne orbiting ▶ Jupiter. Its distance to Jupiter is
experiments for a duration of a few months. 670,900 km, or about 9 Jovian radii. With a
EURECA was released and retrieved by diameter of 3,140 km, Europa is slightly smaller
▶ NASA’s space shuttle. It was the first satellite than our Moon. Its density is 3.0 g/cm3, typical of
designed specifically for microgravity experi- a mixture of rocks including some ice. Its high
ments, providing 105 g conditions for long albedo (0.67) suggests that its surface is mostly
periods. water ice. The surface temperature of Europa
An important feature was reusability: ranges between 125 K at the equator and
EURECA was built to survive during five flights 50 K at the poles.
over a 10-year period. However, it was used only
once, due to lack of funding.
EURECA-1 and its 15 experiments were Overview
launched with shuttle mission STS-46 in July
1992 and were recovered during shuttle mission Europa’s exploration started with the Voyager
STS-57 in June 1993. Among those experiments 2 encounter in 1979. The Voyager camera
Europa 761
revealed a strange surface topography of

low-relief ridges and chaotic terrains with few
impact craters. The pattern of the ridges seemed
to indicate that the icy crust is detached from the
interior and slightly moves with respect to
it. Because Europa is subject to tidal heating due
to the proximity of Jupiter (at a level of about
one-tenth of that of ▶ Io, the Galilean satellite
closest to Jupiter), it was proposed that the icy
crust of the surface might cover a liquid or vis-
E
cous water reservoir. Europa then became a prime
target of the Galileo mission that approached
Jupiter in 1995 and explored the Jovian system
until 2003. Information was retrieved about its
surface features, its internal structure, and its
interaction with the Jovian magnetosphere.
Surface Geology
The most striking features of Europa’s surface are
the series of dark streaks, called “linae” that
crisscross the whole globe. They are innumera-
ble, overlapping up to 1,000 km long and 1 km Europa, Fig. 1 This color image of Europa was taken by
deep, but only hundreds of meters high. In many Voyager 2 during its close encounter on July 9, 1979. The
cases, the ridges are double, often with dark outer array of streaks shows that the surface has been fractured
edges and a central band. Images show that on and filled in some places with material coming from the
interior. The relative absence of features and craters indi-
each side of the lines, the edges have moved cates that the surface is young (# NASA)
relative to each other. These linae are interpreted
as a result of eruptions of warm water, in a sce-
nario similar to the present-day mid-oceanic be estimated on the basis of the number of comets
ridges on Earth. In addition, the fracture patterns and asteroids in Jupiter’s environment, since
on Europa’s surface are different from those these objects would be potential impactors on
expected in the case of an entirely solid planet. the surface. This estimate leads to a surface age
The old and young fractures show different ori- of about 100–300 Ma, which is very young when
entations, the older with the more diverse orien- compared to the whole history of the satellite.
tations. This pattern suggests that Europa’s Most likely, Europa is still geologically active
surface rotates slightly faster than the subsurface today.
material, which implies a liquid or viscous sub-
surface (Fig. 1). Internal Structure and Subsurface Ocean
Europa’s surface also shows fractured and It is now generally accepted that a water ocean
chaotic terrain called “lenticulae,” with diameters lies below the icy crust of Europa. This ocean
ranging from 10 to 100 km. Many are domes that does provide the best explanation for the features
seem to have been pushed up from below. These observed on the surface: linae, lenticulae domes,
domes might have been formed by warm water and fracture patterns. In 2000, another discovery
rising between the cold ices of the outer crust, in provided support to the Europa ocean hypothesis:
a scenario recalling the magma chambers on the magnetometer of Galileo detected a weak
Earth. magnetic moment, induced by the varying part
Impact craters are relatively few, indicating of the Jovian magnetic field. The existence of this
that the surface of Europa is young. Its age can induced magnetic field requires the presence of
762 Europa
highly electrically conducting material inside the depth is beyond our present capabilities, this
satellite. The most plausible candidate is a large result, if confirmed, would have significant impli-
subsurface layer of liquid salt water. The pres- cations for astrobiology.
ence of salts is suspected from the spectral anal-
ysis of the dark surface features; magnesium Interaction with Jupiter’s Magnetosphere
sulfate and sulfuric acids are possible candidates; In 1995, Hubble Space Telescope observations
sulfur components could account for the reddish revealed that Europa has a very tenuous atmo-
color of the streaks. sphere mostly composed of molecular oxygen
What is the internal energy that keeps the O2, with a surface pressure of about 1012 bar.
water warm? Radioactive heating would not be Two years later, the Galileo spacecraft identified
sufficient, as the volume-to-radius ratio is much a tenuous ionosphere resulting from the interac-
smaller than on Earth. In contrast, tidal heating is tion of Europa’s surface with high-energy parti-
the most plausible energy source. Europa is cles of the Jovian ▶ magnetosphere. This
known to be strongly differentiated with a central magnetosphere carries a dense plasma of ener-
core of iron and iron sulfide, surrounded by a getic sulfur and oxygen ions that have escaped
silicate mantle and then by water. Without tidal from Io. The trailing hemisphere of Europa is
effects, water would be in solid state. However, especially exposed to this effect, as shown by
Europa elliptically orbits Jupiter and is in orbital the darkening of its ultraviolet reflectivity. The
resonance with Io and ▶ Ganymede; conse- colliding ions eject water molecules from
quently, although it is in synchronous rotation Europa’s surface (this is the “sputtering effect”);
with Jupiter (showing always the same face to as some of them leave to space, the surface is
Jupiter), the tidal bulge raised by Jupiter changes slowly eroded: the erosion rate is estimated at
slightly in size and orientation as Europa moves about 5–20 cm per million years. This provides
along its orbit. As a result, energy is dissipated additional evidence that the surface of Europa is
inside Europa and models indicate that the max- relatively young. The surface sputtering is also
imum dissipation takes place in the lower part of responsible for the molecular oxygen in the ten-
the ice layer. In 2008, scientists proposed that uous atmosphere. O2 is the major atmospheric
another energy source might come from large component because it does not freeze at the equa-
planetary tides, generated by Rossby waves, due tor. Hydrogen peroxide (H2O2) ice has also been
to the nonzero obliquity of the satellite. detected on Europa by the Near Infrared Mapping
What could be the thickness of the ice crust? Spectrometer of Galileo, and hydrogen peroxide
According to the models, it could range between acid mixtures might account for some features of
a few kilometers and a few tens of kilometers. the surface spectrum. Molecular hydrogen cannot
The study of Europa’s largest craters, surrounded remain on Europa’s surface because the satel-
by concentric rings and filled with flat and fresh lite’s gravity field is too low. It escapes in a
ice, suggests a 10–30 km thick ice crust, includ- torus, along with atomic and molecular oxygen,
ing a ductile warm ice layer, over a liquid ocean to form a neutral torus along the orbit of Europa.
that might be about 100 km deep. According to This torus has been identified by the Galileo and
models of internal structure, the liquid ocean is Cassini spacecraft.
expected to be in direct contact with the silicate
mantle. Potential for Extraterrestrial Life
An important result has come recently from Europa’s interior is considered as a possible can-
the reanalysis of Galileo images of chaos terrains didate for extraterrestrial life in the solar system.
by a group of scientists from the University of Indeed, there are three conditions that seem to be
Texas at Austin. This work suggests that these required for the development of life on Earth: the
areas are formed above liquid water lenses presence of carbon, an energy source, and liquid
located within an ice shell as shallow as 3 km. water. Many carbon-rich compounds have been
Although drilling the surface of Europa at 3 km found in comets and meteorites, and such
Europa Analogues 763
elements must have been brought to Europa by References and Further Reading
meteoritic impacts. Even if impact craters have
been removed by resurfacing, Europa underwent Hall DT et al (2006) Detection of an oxygen atmosphere
on Jupiter’s moon Europa. Nature 373:677–679
an intense meteoritic bombardment like that on
Hurford TA, Sarid AR, Greenberg RN (2007) Cycloidal
▶ Ganymede and ▶ Callisto. The energy source, cracks on Europa: improved modelling and non-
as mentioned above, is provided by tidal heating synchronous rotation implications. Icarus 186:218–233
that comes in addition to radioactive heating. Life Kivelson MG et al (2000) Galileo magnetometer measure-
ments: a strong case for a sub-surface ocean at Europa.
could exist in an under-ice ocean, in an environ-
Science 289:1340–1343
ment comparable to the Earth’s hydrothermal McCord TB et al (1998) Salts on Europa’s surface
vents, or the Antarctic Lake Vostok. On Europa, detected by Galileo’s near infrared mapping spectrom-
eter. Science 280:1242–1245
E
free radicals delivered by subduction of surface ice
Pappalardo RT, McKinnon WB, Khurana KK
may provide a sufficient source of free oxygen,
(2009) Europa. University of Arizona, Tucson, p 727
even in the absence of photosynthesis. Life on Sotin C, Head JW, Tobie G (2002) Europa: tidal heating of
Europa, if it exists, could be clustered around upwelling thermal plumes and the origin of lenticulae
hydrothermal vents on the ocean floor or below and chaos melting. Geophys Res Lett 29(8):1233.
doi:10.1029/2001GL013844
the ocean floor. Alternatively, it could exist cling-
Tyler RH (2008) Strong ocean tidal flows and heating on
ing to the lower surface of Europa’s ice layer, moons of the outer planets. Nature 456:770–772
much like algae and bacteria in Earth’s polar
regions, or float freely in Europa’s ocean. If
Europa’s ocean is too cold or too salty, only
▶ extremophiles could survive in its environment. Europa Analogues
It should be remembered that there is not yet
any evidence that life actually exists on Europa. Felipe Gomez
However, there is a large consensus in the scien- Centro de Astrobiologı́a (CSIC/INTA), Instituto
tific community about the existence of an ocean Nacional de Técnica Aeroespacial, Torrejón de
of salty liquid water, probably in contact with a Ardoz, Madrid, Spain
silicate mantle, under an icy crust which is prob-
ably several kilometers or several tens of kilome-
ters thick. The fact that Europa is a potential Keywords
candidate for extraterrestrial life has motivated
the scientific community to design an ambitious Antarctica; Jupiter; Europa
mission, JUICE (Jupiter and Icy moons
Explorer), which has been selected by ESA for a
launch in 2022 and an arrival at the Jupiter system Definition
in 2030. The spacecraft will achieve several
flybys of Jupiter and the Galilean satellites, Extreme field sites on Earth that mimic some of the
including two flybys of Europa, and will be put conditions on Jupiter’s moon Europa are consid-
in orbit around Ganymede in 2033 for an in-depth ered Europa analogues. Some Earth natural field
exploration of the habitability conditions of the sites are extreme, due to very low temperature,
icy moons. high salinity, or high pressure conditions. These
extreme conditions partially characterize the envi-
ronment of Europa. Lake Vostok in Antarctica is
See Also one of the best recognized Europa analogues. Due
to the thick ice crust, a liquid water reservoir
▶ Callisto underneath is stable due to salinity and high pres-
▶ Cryovolcanism sure conditions. Lake Tirez in La Mancha (Spain)
▶ Ganymede is considered a geochemical analogue of Europa
▶ Io due to the ionic content of its waters.
764 European Space Agency
See Also ▶ Gaia Mission

▶ Europa ▶ International Space Station
▶ Vostok, Subglacial Lake ▶ Mars Express
▶ PLATO 2.0 Satellite
European Space Agency
David W. Latham European Space Exposure Facility

Harvard-Smithsonian Center for Astrophysics, (ESEF)
Cambridge, MA, USA
▶ COMET (Experiment)
Definition
The European Space Agency (ESA) was

Euryarchaeota
established in 1975 and is responsible for coordi-
nating the exploration of space by its member
Ricardo Amils
countries, which numbered 18 in 2010: Austria,
Belgium, Czech Republic, Denmark, Finland,
Universidad Autónoma de Madrid, Madrid, Spain
France, Germany, Greece, Ireland, Italy, Luxem-
bourg, Netherlands, Norway, Portugal, Spain,
Sweden, Switzerland, and United Kingdom and
Definition
Canada as an associated member. ESA’s program
includes human space flight, primarily through
Euryarchaeota is one of the four phyla of the
participation in the ▶ International Space Station,
domain ▶ Archaea. Euryarchaeota comprises a
and a variety of scientific missions. Of particular
physiologically diverse group of Archaea: all
interest to astrobiology is Gaia, an all-sky astro-
known methanogens, extreme halophilic
metric survey mission that will detect hundreds,
Archaea, ▶ hyperthermophiles such as
if not thousands, of extrasolar planets. ESA is
Thermococcus and Pyrococcus, most
also committed to the exploration of Mars, most
acidophilic-thermophilic prokaryotes including
notably with the successful Mars Express Mis-
Picrophilus, and the thermophilic-acidophilic
sion launched in 2003, and the exploration of
cell wall-less Thermoplasma. A large number of
▶ Titan with the successful landing at the surface
Euryarchaeota produce methane (CH4) as an inte-
of the satellite of the ▶ Huygens probe in 2005,
gral part of their metabolism (see
and in the exploration of comets with the Giotto
▶ methanogens). It is considered a very ancient
mission in 1986 and the ongoing Rosetta mission.
metabolism due to the unique set of enzymes
involved. The extreme halophilic Archaea are a
diverse group of Euryarchaea that inhabit highly
See Also saline environments (see ▶ halophile). Most spe-
cies require 2–4 molar salt concentration for opti-
▶ Astrometric Planets mal growth, and they are able to grow in the
▶ Beagle 2 presence of saturating salt conditions.
▶ Cassini-Huygens Space Mission Picrophilus is capable of growing below pH 0.
▶ CoRoT Satellite Pyrococcus, which literally means fireball, has an
▶ Exoplanets, Discovery optimal growth at 100 C.
Evaporites, Archean 765
See Also of evaporite minerals on planetary surfaces may

record the past existence of water. The ▶ Mars
▶ Acidophile Rover Opportunity has discovered the presence
▶ Archaea of gypsum and ▶ jarosite in the Burns Formation,
▶ Halophile Meridiani Planum, Mars, which are interpreted
▶ Halotolerance as deposits formed by the evaporation of acidic
▶ Hyperthermophile waters.
▶ Methanogens
See Also
E
▶ Carbonate
EUV ▶ Jarosite
▶ Mars
▶ Extreme Ultraviolet Light ▶ Mars Exploration Rovers
▶ Natron
▶ Soda Lakes
▶ Thermonatrite
Euxinic Ocean ▶ Trona
▶ Sulfidic Oceans
Evaporites, Archean
Evaporite Sami Nabhan

Institute of Geological Sciences, Freie
Daniele L. Pinti Universität Berlin, Berlin, Germany
Montréal, QC, Canada Keywords
Evaporites; Barite; Gypsum; Carbonate; Chert;

Synonyms BIF; Archean; Continental growth
Evaporitic rocks and minerals

Definition
Definition Evaporites are chemical sediments and sedimen-

tary rocks that form as result of an increasing
Evaporites are a class of sedimentary rocks and concentration of ions in water as precipitates
minerals precipitated from evaporating aqueous either at the surface or at the base of the water
fluids. Common evaporitic minerals from waters body. The majority of evaporitic sediments and
modified from the composition of the present-day sedimentary rocks on Earth consists of carbon-
ocean include calcite (CaCO3), dolomite (CaMg ates, sulfates, or halides. Archean evaporites are,
(CO3)2), anhydrite (CaSO4), gypsum (CaSO4 to their largest part, either of a restricted-marine
2H2O), and halite (NaCl), which precipitate out facies or hydrothermally related. Open-marine
of solution in the reverse order of their solubil- evaporites are completely absent, terrestrial
ities from evaporating seawater. The occurrence evaporites nearly so.
766 Evaporites, Archean
Overview platform in Ontario, Canada. At about 2.6 Ga,

platform-type carbonates became established
Evaporites compose a disproportionally small (Grotzinger and Kasting 1993; Harnmeijer
percentage of the Archean rock record because 2010), substantially composed of original arago-
their preservation potential is generally low and nite (now mainly dolomitized) and exemplified
because conditions for their formation may have by the ~2.55-Ga-old Campbellrand-Malmani car-
been poor due to small and few ▶ continents, bonate platform with individual beds of up to
lower continental freeboard, a perceived hot and 65 % encrusting carbonate. This suggests that
humid climate, and an unknown saturation state seawater was supersaturated with respect to ara-
of the oceans. Ocean salinity and surface water gonite and calcite and that its pH was neutral to
temperature are major controlling factors for the slightly alkaline (Sumner and Grotzinger 2004).
precipitation of evaporites. Estimates of the In some parts of the Archean carbonate plat-
Archean ocean temperature based on oxygen iso- forms, the presence of pseudomorphs of carbon-
tope composition of early diagenetic ▶ chert ates and chert after halite crystals combined with
units are debated, as is the salinity of Archean an absence of ▶ sulfate pseudomorphs within
ocean water. Archean ocean temperatures, as supratidal lithofacies zones (Boulter and Glover
indicated by the isotope composition of Archean 1986; Sumner and Grotzinger 2004; Eriksson
cherts, were approx. 55–85 C (Knauth 2005). et al. 2005; Gandin et al. 2005) has been
Estimates of Archean ocean water salinity based interpreted to either reflect a generally low sulfate
on “adding back” the volume of halite in the content or a low calcium content of seawater, a
geological record to the oceans would result in a consequence of extensive carbonate precipitation
salinity of 1.2–2 times the modern value (Knauth (Boulter and Glover 1986; Grotzinger and
2005). This number, however, is highly depen- Kasting 1993; Sumner and Grotzinger 2004;
dent on estimates of the size of subsurface salt Eriksson et al. 2005). On the other hand, pseudo-
deposits and of hypersaline brines in sedimentary morphs after swallowtail gypsum and enterolithic
basins. Other factors controlling evaporation gypsum have been described by Gandin
include the ionic composition of Archean ocean et al. (2005). Some of the crystal fans interpreted
water and its pH value. The suggestion of an early as aragonite fans resemble gypsum domes in their
ocean saturated in sodium carbonate (the “soda structure (Hardie 2003; Gandin et al. 2005).
ocean”) would result in a pH of ~10 (Kempe and Calcitization of former sulfate evaporites is
Degens 1985; Gargaud et al. 2005). The presence interpreted as result of bacterial sulfate reduction
of reduced iron (Fe2+) in the Archean ocean, and the organic diagenesis of pyrite (Gandin
however, indicates a pH <6 (Gargaud et al. 2005). Such a metabolic process would
et al. 2005). constitute a second link to widespread microbial
Archean evaporites are commonly preserved life in the Archean, aside from the abundant mor-
as pseudomorphs and silicified from their precur- phological evidence provided by stromatolites in
sors (gypsum, nahcolite, and aragonite). The carbonate platforms.
majority of evaporites are preserved as Sulfate deposits formed as thinly bedded bar-
dolostones and limestone that originally precipi- ite of early and mid-Archean age are well
tated as carbonate cements such as aragonite and represented in the ~3.3-Ga-old Sargur Group,
magnesian calcite (Grotzinger and Kasting 1993) ▶ Dharwar Craton, central India (Jewell 2000);
or as ▶ stromatolites, a common component of the ~3.45-Ga-old Onverwacht and the ~3.24-Ga-
Archean carbonate platforms (Grotzinger and old Fig Tree Groups of the ▶ Barberton Green-
Knoll 1999). Most carbonates older than ~3 Ga stone Belt, South Africa (Lowe and Fisher
are of peritidal sedimentary facies and found in Worrell 1999); and the 3.45-Ga-old North Pole
▶ greenstone belts of South Africa, Australia, chert-barite unit of the Warrawoona Group,
and India (Harnmeijer 2010). The oldest known Western Australia (Jewell 2000). All of them
carbonate platform is the ~2.9-Ga-old Steep Rock are of marine origin and show primary
Evaporites, Archean 767
sedimentary and partly diagenetic structures depending on ocean water chemistry (Huston and
(Buick and Dunlop 1990; Jewell 2000). For Logan 2004; Pufahl and Hiatt 2012). They are
some deposits, their possible origin by volcanic- interpreted either as deepwater precipitates or as
hydrothermal exhalation is discussed (Reimer precipitates developed during times of restricted
1980). Parts of the North Pole barite have been clastic supply.
interpreted as pseudomorphosed gypsum, based
on interfacial angles of individual crystals (Buick
and Dunlop 1990; Jewell 2000; Shen et al. 2001). Basic Methodology
▶ Sulfur isotope variations within these sulfates
in microcrystalline ▶ pyrite suggest the existence Field observations (mapping, documentation of
E
of sulfate-reducing bacteria (Shen et al. 2001; sedimentary structures, construction of strati-
Shen and Buick 2004). graphic cross sections)
Overall, the presence of evaporitic sulfates in Thin-section microscopy
the 3.5–3.2 Ga Archean rock record may indicate Analysis of major and trace element distribution
a global sulfate-bearing, partly oxidized, and Raman spectroscopy for fluid and gas inclusions
stratified ocean (Huston and Logan 2004). Aside Stable isotope fractionation (S, C, O, N, Sr, Fe)
from the three deposits mentioned above, only
two additional bedded sulfate deposits are
Applications
known in the geologic record prior to the GOE
at 2.4 Ga, after which they become more abun-
Evaporites are sedimentary rocks that reflect
dant (Huston and Logan 2004), possibly due to
atmospheric and hydrospheric conditions during
low concentrations of sulfate in the Archean
the time of their formation. Therefore Archean
ocean (Boulter and Glover 1986; Grotzinger and
evaporites can be used to constrain Archean
Kasting 1993; Sumner and Grotzinger 2004;
atmospheric and seawater composition. Terres-
Eriksson et al. 2005; Gargaud et al. 2005). How-
trial evaporites can also be used to provide infor-
ever, a partly oxidized ocean may not be required
mation on Archean groundwater and to infer
to produce low seawater sulfate from volcanic
climatic conditions. Evaporites are often related
SO2 because SO2 reacts with H2O to H2S and
to or aided by biogenic processes and may thus
H2SO4 even in the absence of oxygen (Huston
preserve traces of early life.
and Logan 2004; Gargaud et al. 2005).
Terrigenous evaporites are exceedingly rare in
the Archean. Nabhan et al. (2013) describe wide-
spread silicified former sulfate concretions from See Also
the vadose zone of a fluvial to supratidal facies of
the ~3.2-Ga-old Moodies Group, South Africa, ▶ Archean Environmental Conditions
and interpreted them to have formed during early ▶ Banded Iron Formation
diagenesis by evaporating groundwater in a semi- ▶ Barite
arid or arid coastal plain. ▶ Campbellrand-Malmani Platform, South
Some Archean cherts and possibly banded- Africa
iron formations (BIF) can be considered evapo- ▶ Chert
rites where the involvement of biologic processes ▶ Evaporite
is uncertain or unlikely. Some cherts may form ▶ Jaspilite
abiogenically in subtidal environments due to ▶ Magnetite
silica oversaturation of pore waters during evap- ▶ Moodies Group
oration (Pufahl and Hiatt 2012). ▶ Banded-iron ▶ North Pole Dome (Pilbara, Western Australia)
formations are composed of a mixture of silica ▶ Oceans, Origin of
with iron-bearing minerals such as hematite, ▶ Precambrian Oceans, Temperature of
magnetite, siderite, and pyrite which precipitate ▶ Pyrite
768 Evaporitic Rocks and Minerals
▶ Stromatolites Nabhan S, Heubeck C, Homann M (2013) Evaporitic sul-

▶ Sulfate Minerals fate concretions, Moodies Group (~3.2 Ga, Barberton
Greenstone Belt, South Africa). Mineral Mag 77:1818
▶ Sulfate Reducers Pufahl PK, Hiatt EE (2012) Oxygenation of the Earth’s
▶ Sulfur Isotopes atmosphere-ocean system: a review of physical and
▶ Warrawoona Group chemical sedimentologic responses. Mar Pet Geol
32:1–20
Reimer TO (1980) Archean sedimentary barite deposits of
the Swaziland Supergroup (Barberton Mountain Land,
References and Further Reading South Africa). Precambrian Res 12:393–410
Shen Y, Buick R (2004) The antiquity of microbial sulfate
Boulter CA, Glover JE (1986) Chert with relict hopper reduction. Earth-Sci Rev 64:243–272
moulds from Rocklea Dome, Pilbara Craton, Western Shen Y, Buick R, Canfield DE (2001) Isotopic evidence
Australia: an Archean halite-bearing evaporite. Geol- for microbial sulphate reduction in the early Archaean
ogy 14:128–131 era. Nature 410:77–81
Buick R, Dunlop JSR (1990) Evaporitic sediments of early Sumner DY, Grotzinger JP (2004) Implications for
Archaean age from the Warrawoona Group, North Neoarchaean ocean chemistry from primary carbonate
Pole, Western Australia. Sedimentology 37:247–277 mineralogy of the Campbellrand-Malmani Platform,
Eriksson KA, Simpson EL, Master S, Henry G (2005) South Africa. Sedimentology 51:1–27
Neoarchaean (c. 2.58 Ga) halite casts: implications
for palaeoceanic chemistry. J Geol Soc 162:789–799
Gandin A, Wright DT, Melezhik V (2005) Vanished evap-
orites and carbonate formation in the Neoarchaean
Kogelbeen and Gamohaan formations of the
Campbellrand Subgroup, South Africa. J Afr Earth Evaporitic Rocks and Minerals
Sci 41:1–23
Gargaud M, Barbier B, Martin H, Reisse J (eds) ▶ Evaporite
(2005) Lectures in astrobiology. Adv Astrobiol
Biogeophysics 1. XXVI:793 pp
Grotzinger JP, Kasting JF (1993) New constrains
on Precambrian ocean composition. J Geol 101:
235–243
Grotzinger JP, Knoll AH (1999) Stromatolites in Precam-
Evo-devo
brian carbonates: evolutionary mileposts or environ-
mental dipsticks? Earth Planet Sci 27:313–358 Stéphane Tirard
Hardie LA (2003) Secular variations in Precambrian sea- Centre François Viète d’Histoire des Sciences et
water chemistry and the timing of Precambrian arago-
nite seas and calcite sea. Geology 31:785–788
Harnmeijer JP (2010) Squeezing blood from a stone: des Techniques de Nantes, Nantes, France
inference into the life and depositional environments.
PhD thesis, University of Washington, Washington
Huston DL, Logan GA (2004) Barite, BIFs and bugs:
evidence for the evolution of the Earth’s early hydro-
Definition
sphere. Earth Planet Sci Lett 220:41–55
Jewell PW (2000) Bedded Barite in the geologic record. “Evo-devo” is the term used to designate the
SEPM Spec Publ 66:147–161 relationship between evolution and developmen-
Kempe S, Degens ET (1985) An early soda ocean? Chem
Geol 53:95–108
tal biology. It corresponds to the evolutionary
Knauth LP (2005) Temperature and salinity history of the developmental biology. This term “evo-devo”
Precambrian ocean: implications for the course of was created in the 1980s, after the discovery of
microbial evolution. Palaeogeogr Palaeoclimatol the genes that regulate development of the animal
Palaeoecol 219:53–69
body, notably the Hox gene complex.
Lowe DR, Fisher Worrell GF (1999) Sedimentology, min-
eralogy, and implications of silicified evaporates in the During the last decades, the expansion, the
Kromberg Formation, Barberton Greenstone Belt, distribution, and the changes in the embryonic
South Africa. In: Lowe DR, Byerly GR (eds) Geologic expression of these genes in taxa have been asso-
evolution of the Barberton Greenstone Belt, South
ciated to explain evolutionary changes in animal
Africa, Geological Society of America Special Paper
329. Geological Society of America, Boulder, organization.
pp 167–187
Evolution of Species, Islamic Ideas 769
literature “written in a highly expressive and

Evolution of Species, Islamic Ideas ornate style. . .with its emphasis on philosophical
and religious edification” rather than a zoological
Salman Hameed treatise. Al-Jahiz believed in spontaneous gener-
Hampshire College, Amherst, MA, USA ation of life, but he commented on the intercon-
nectedness of life and believed that every plant
and animal had its place in the scheme designed
Keywords by God. It is in this context that he described
ecosystems and commented on adaptation of spe-
Evolution; The great chain of beings; Al-Jahiz; cies, where stronger animals “feed upon weaker
E
Brethren of Purity; Ibn-Miskawayh species than itself ” and that “each animal is the
food for a species that is stronger than itself”
(Stott 2013). The works of Ibn-Miskawayh
Definition (940–1030 C.E.) and a group of tenth-century
pseudonymous writers in Basra known as
There are few medieval Islamic scholars who ▶ Ikhwan al-Safa (Brethren of Purity) drew on
speculated on the evolution of species (e.g., the idea of the Great Chain of Being that
Al-Jahiz, Ibn-Miskawayh, and a group of writers connected all life (and also minerals, with alum
using the pen name the Brethren of Purity). In and vitriol being the lowest types) in a hierarchi-
most instances they believed in the creation of cal structure, but then speculated on the possibil-
species by God and a hierarchy of God’s creation ity of transformation of species within the ladder.
from minerals, plants, and animals to humans, While apes are mentioned to be close to humans,
angels, saints, and prophets. Al-Jahiz, however, their hierarchical structure also includes angels,
did comment on the interconnectedness of life saints, and prophets, thus making it hard to argue
and described an ecosystem based on the adapta- for a straightforward theory of evolution
tion of species to local environment, but the (Elshakry 2014). The Ikhwan are considered
Brethren of Purity considered adaptation as a keen observers of the biological world, but con-
gift from God rather than a process responding sider adaptation as God’s grace to the species
to environmental conditions. rather than a process. Similarly, while they do
identify some animals as intermediate species
(e.g., they call the giraffe a cross between a
Overview camel and an ass), they still see these as stable
forms of its kind (Goodman and McGregor
There are diverse claims about medieval Islamic 2009). Much of the uncertainty in the interpreta-
ideas on the evolution of species and there is tions comes from the fact that the terms used in
limited scholarship available on this medieval literature are “change,” “stage,” and
topic. Some of the uncertainty lies in the late “transformation” and scholars have to interpret
nineteenth- and twentieth-century translations this in light of the use of “evolution” (Guessoum
and interpretations of medieval works in light of 2011).
Darwin’s theory. Sarton (1927) and Bayrakdar
(1983) made strong claims that Al-Jahiz
(776–886 C.E.) in his book, The Book of Animals
(Kitab al-Hayawan), came close to natural selec- Bayrakdar M (1983) Al-Jahiz and the rise of biological
tion and some form of a theory of evolution. evolutionism. Islam Q 21:149–155
Egerton (2002) disputes the strong claim but Egerton FN (2002) A history of the ecological sciences,
part 6: Arabic language science: origins and zoological
attributes the origin of a detailed idea of a food
writings. Bull Ecol Soc Am 83(2):142–146
chain to Al-Jahiz. According to Elshakry (2014), Elshakry M (2014) Reading Darwin in Arabic,
The Book of Animals was more a work of 1860–1950. The University of Chicago Press, Chicago
770 Evolution, Biological
Goodman LE, McGregor R (2009) The Brethren of Purity: Biology. Evolution is sometimes understood as a
the case of animals versus man before the King of the movie scenario, a big story to tell. Evolution is also
Jinn: an Arabic critical edition and English translation
of epistle 22. Oxford University Press, New York seen, in our imaginations, as a big tree: the tree of
Guessoum N (2011) Islam’s quantum question: reconcil- ▶ life. Evolution is also more or less understood as
ing Muslim tradition and modern science. I. B Tauris, a progress. Evolution is seen as a process by which
London living things do change over time. Last but not
Sarton G (1927) An introduction to the history of science,
vol 1. Williams and Wilkins Press, Baltimore least, evolution is meant as the general theory of
Stott R (2013) Darwin’s ghosts: the secret history of Biology, Palaeontology, and Anthropology. The
evolution. Spiegel & Grau Publications, New York two meanings of the word that scientists would
like to retain are the last two. In the eighteenth
century, evolution (from latin evolutio, from the
Evolution, Biological verb evolvere) originally meant the deployment of
an inner content, and was employed as we use
Guillaume Lecointre and Pierre-Henri Gouyon today embryonic development. The famous geol-
Département Systématique et Evolution, UMR ogist, Charles Lyell, used the term in the sense of
7138 CNRS-MNHN-UPMC-IRD, Muséum transformation of ▶ species for the first time in the
National d’Histoire Naturelle, Paris Cedex 05, second volume of his Principles of Geology
France (1832). Herbert Spencer (1862) popularized the
term in a very wide sense including transformation
of species through times (Tort 1996). Charles Dar-
Keywords win did not use the word in the first editions of his
famous book The Origin of Species (Darwin
Adaptation; Classification; Drift; Facts; History; 1859), but did in the fifth and sixth editions.
Mutualism; Natural selection; Phylogeny; Sexual However, in a more general context, evolution
selection; Species; Systematics; Theory; is a term used to designate a change. Evolution of
Variation the political situation of a country, evolution of the
social condition of workers, evolution of the health
of a patient following a treatment against a given
Synonyms sickness, evolution of the weather, evolution of the
universe, etc., are examples of common uses of the
Biological evolution term. The question of transferring the term “evolu-
tion” to designate precise phenomena in chemistry
or astronomy is meaningless if the term is under-
Definition stood in that general sense of “change.” The whole
physical world is, obviously, subject to change.
In Biology, evolution is the change of organisms The question becomes more interesting if, by “evo-
through their variation, then transmission of these lution,” a particular process of change is meant.
variations to their progeny (whatever the way Evolution would be “change” through heritable
they are transmitted), and, eventually, ▶ selec- variation of physical entities and selection of
tion of some phenotypic variant in the long run some variants by the conditions of the environ-
as a result of environmental conditions. ment – whether biological, chemical, or physical.
Evolution: What’s in a Word? Evolution in Biology: Theory and Facts

When speaking about evolution people can have Evolution is the general theory of Biology,
different meanings in mind even within the field of Palaeontology, and Anthropology. The role of a
Evolution, Biological 771
theory is to coordinate facts, laws, models, and This is not true. Two epistemological sides of
hypotheses, and to maximize the global consis- evolutionary sciences must be distinguished here.
tency of the whole. A theory explains facts and First of all, evolution is supported by many
allows us to predict some yet unobserved facts. experimental proofs. Organisms with short gen-
When a theory is corroborated, as highly as pos- eration times are grown in the lab (fungi, bacteria,
sible, it is still a theory. Unfortunately, scientists fruit flies, etc.,), variation can be measured, and,
stop speaking about their favorite theories as knowing parameters of selective pressures,
“theories” when their reliability is maximal. But changes of variant frequencies in populations
there’s no epistemological change to justify this. can be measured and predicted. Famous exam-
As a result the public has a kind of implicit ples of experiments in evolutionary sciences of
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hierarchy in mind: a theory is dubious, facts are the twentieth century involved the fruit fly Dro-
serious. But evolution is still a good theory and its sophila melanogaster or the fungi Ascomyces.
role as a theory has not changed since Darwin. Its Today, it is possible to make bacteria like
degree of corroboration is today as high as the Escherichia coli perform protein engineering
theory of chromosomal heredity or the theory of just by introducing selective pressures at the lab.
continental drift. Hierarchy distinguishing theo- Hypothetico-deductive reasoning is at work: if
ries from facts is nonsense for Science in general. the premises and the law (or the rule) are true,
Theories are useful to organize facts, even to the result is true. Evolutionary mechanisms are
simply speak about facts. Everyone should therefore either experimented/reproduced at the
remember that cars, planes, fridges, or phones lab or observed in the field, and can lead to short
are built according to theories and not facts. term predictions. However, products of evolution
There are evolutionary facts, like fossil discover- in the wild are contingent. Each population or
ies or DNA sequences, and at the same time there each species is produced by laws of mutations,
is a very good theory: evolutionary theory. recombinations, genetic drift, and ▶ natural
A theory without facts is fantasy; facts without a selection, but the result is always historically
theory are chaos. Such an embedment of facts singular and cannot be predicted in the long
within theories leads to the question of theoretical term. One of the reasons is that all parameters of
changes. If facts can only make sense through a the environment undergo changes in a contingent
given theory, how can we change that theory for manner. This is why evolutionary sciences are
an alternate one? Usually a given fact is evaluated also historical sciences.
within (at least) two competing theories. The Secondly, evolution is supported by historical
parsimony of a theory is the same measurement proofs, which are another kind of scientific
as the coherence of facts (or global consistency): proofs. If the premises are true, the result is only
the more coherent is a set of facts the less ad hoc probable. We have a number of signs and facts.
hypotheses we need to add to maintain the theory. We create a scenario that maximizes consistency
This is why we prefer, among the competing of the whole set of facts, minimizing ad hoc
theories, the most parsimonious theories. hypotheses. This is the reasoning of historians,
Claiming that evolution “is only a theory” is policemen, and phylogeneticists. The reasoning
therefore meaningless for scientists. is called abductive: we infer what might have
occurred in the past to explain the state of facts
Evolution Through Multiple Sciences in the present. The scenario is made to explain the
Sometimes scientists are asked to provide “evi- present facts and signs. Abduction is a “reverse
dence” for evolution. Evolution being understood deduction,” deduction being found in a situation
as a scenario, they are expected to come back from where present facts and signs would explain the
the past with pictures and films captured with their scenario. By using algorithms that maximize con-
camera. Speakers in creationist or intelligent sistency, phylogeneticists do produce statements
design books, websites, or “museums” claim that that can be tested, i.e., reproduced or refuted,
scientists actually know nothing from the past. even if the reasoning is abductive.
Key Research Findings (Mus musculus) to the Madeira Islands with

their ships. These islands have high mountains
Evolution: Biological Processes that separate valleys. Mice settled in the valleys
The general process of biological evolution is in small populations, almost isolated from one
based on variation. Among a set of individuals another. Chromosomal changes did occur. Some
that are able to interbreed, one can notice slight of the resulting chromosomal formulae reached a
differences. In the hall of the airport, at a given frequency of 100 % in one valley just by drift.
time no two people have the same face. This is Those formulae, once analyzed, show that mice
basically due to the fact that living matter is from one valley cannot breed with mice from the
constantly altered. Individuals belonging to the neighboring valley anymore. Reproduction is
same species are not identical, and they can differ impossible because, once fecundation has
and vary in traits that are inheritable. Indeed some occurred, genetic material cannot be harmoni-
of the variations can be transmitted to the off- ously distributed among daughter cells.
spring. These varying traits are those involved in Populations changed, and new species (as we
evolution. Different versions of those traits can conventionally limit species with obstacles in
be present in a species. Each version is (or may the interbreeding) appeared within the past five
be) represented by several individuals. Some centuries.
individuals will have a high number of descen- Charles R. Darwin (1809–1882) introduced
dants, others only a few, others none at all, either the notion of natural selection. Among versions
because the variations influence the reproductive of a given trait, some of them confer to the indi-
success (selection) or just by chance (drift). We vidual that bears them a certain advantage in
shall now examine these two processes. When the terms of number of offspring. The version of the
variation does not influence the reproductive out- trait is advantageous within a certain environ-
put of individuals, there is no selection. In this ment. If the trait is heritable and if the environ-
case, each generation represents a sampling ment is stable enough for a while, the version of
among the hereditary diversity of the preceding the trait can easily reach a frequency of 100 % in
one. As a result the frequency of a given version the species. This tendency to accumulate traits
of the trait does vary with time. If the global favored by a given environmental condition is
number of individuals is small, these fluctuations called ▶ adaptation. The process is acting on
can have a high effect on the relative proportions many traits at the same time, as organisms are
of the versions within the species. This is called a made of multiple traits. The differential repro-
“drift” in frequencies of versions. Eventually, a ductive success among versions of a trait
given version will reach a frequency of 100 % or (disappearing of disadvantageous versions and
disappear just by chance. The smaller the number increase of copies of advantageous versions) is
of individuals that can give offspring, the faster called natural selection. Note that many versions
drift is (a sample differs more from the original are just neutral with regard to the environment:
set when it is small). The species has thus neither advantageous nor disadvantageous. Note
changed. If the number of individuals is high, that drift can fix slightly disadvantageous traits
drift is very slow and mutations which introduce just by chance. Note that selection of a strongly
variation compensates for the loss of diversity. advantageous trait can have side effects, i.e., be
This process constantly maintains a number of accompanied with disadvantageous conse-
different versions within the species – this is quences onto another trait. There is no “perfect”
called polymorphism. We must keep in mind organism: natural selection is a permanent trade-
that once a version has disappeared, or once a off among traits that are not fully independent
version has reached a frequency of 100 %, a new from each other. For instance, aggressiveness is
version of the same trait can appear again: life is very important among female spotted hyenas.
constantly altered. For example, five centuries The more aggressive they are, the more they eat,
ago the Portuguese brought domestic mice the more their daughter eats, and the higher their
social rank, which is transmitted to daughters (not a complex process and minor changes in the
to sons). Males are smaller and outside this com- timing or functioning of each step can have
petition. Cubs are aggressive right after their important consequences on the shape of the
birth. They fight with each other until death. future individual. These variations, occurring
Young females keep being aggressive all through independently from the future needs of the organ-
their lives. Their brain had been marked by high ism but sorted out through the relative success
levels of testosterone in the very early period of (in terms of survival and reproduction) of the
their development. Indeed the placenta of spotted individuals issued from them (i.e., natural selec-
hyenas has enzymes that transform the circulat- tion), explain the evolution of the shape of organ-
ing estradiol of the mother into testosterone that isms and constitute the “evolutionary tinkering.”
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circulates in the blood of the fetus. The young Natural selection has many facets: the ways by
maintain high levels of testosterone in utero which a given version can have a higher number
which has the advantageous consequence of of descendents are many. Let’s call them modal-
early aggressiveness, but also the disadvanta- ities and let’s list five of them. The first modality
geous side effect of having the female external of natural selection is competition among indi-
sexual organs being deviated towards a male viduals of the same species for resources (food,
development. As a result adult females have a home). This is the first one we think about
peniform clitoris, an obturated vagina, a pseudo- because Darwin himself emphasized the “strug-
scrotum. They give birth through the clitoris, gle for life” in his main book, an idea coming
which is a unique case among mammals. Because from the simple comparison between the increase
of this anatomical curiosity they lose 60 % of of resources and the potential increase of off-
their first born progeny: the umbilical duct is spring of the species exploiting them. The second
shorter than the corridor they must go through, increase is exponential, the first one is arithmetic:
provoking the separation of the placenta before all individuals cannot get the resources they need.
the cub is out, which sometimes dies from anoxia. In other words, the populating power of a species
This gives an example – among so many is by far higher than the actual state of
others – that organisms are far from perfect and populations: if every elephant could have all the
then not “designed.” food it needs, within a few decades Africa would
Perfection is not a scientific notion anyway, be all covered with elephants. There must be a
but nested within the realm of individual values. competition somewhere, leading to the fact that
François Jacob has compared evolution by natu- any inheritable version of a trait that favors the
ral selection to tinkering. Contrarily to an engi- ability of individuals to get more food must have
neer who designs the final product he wants to led to more offspring against versions that did not
achieve and produces the pieces in order to reach confer such an advantage. However, competition
this goal, evolution transforms existing parts for food is by far not the only modality.
through random variation and just sorts out, A second modality is competition among indi-
through selection, those which are favored within viduals of the same sex for sexual intercourse,
a given context. This is as if planes had been built called sexual selection. Any change of traits
by transforming a car. In evolution, tinkering can favoring sexual attraction, if heritable, must
be seen at work in different cases. A classic have led to more offspring and must have been
example is the feathers of birds, originally rapidly spread through the species. Darwin wrote
selected for their thermal properties, then used an entire book about sexual selection (Darwin
for flight. One can also cite the fact that the same 1871). It is clear that there is often a trade-off
organ can play the role of swim bladder in some between natural selection and sexual selection.
fishes or of lungs in other animals, with some A classical example is the magnificent caudal
intermediate forms perhaps used at both roles. feathers of the male of the peacock. Observations
Any new form is a result of a transformation of and experiments led to the conclusion that such
a preceding one. The development of embryos is an apparatus has a real impact on females’
preferences: the bigger the feathers, the higher as the association of individuals from different
the chance for the male to be chosen as a sexual species having both more offspring when associ-
partner. However, the whole bulk of caudal ated than when separated. Any change that could
feathers is a very long and embarrassing burden favor such phenomena must have been spread
that does not allow the male to escape rapidly through generations.
from predators. Such a characteristic is thus the A fifth modality of natural selection is partner-
result of a compromise between the advantage in ship within the species. This could explain the
seducing females on one side and the disadvan- origin of multicellularity. Traits are transmitted
tage in escaping from predators in the given to the progeny – to a wide extend – through the
environment. transmission of genetic material. Any phenotypic
The third modality of natural selection is trait inducing a behavior which increases the
linked to predation. The more we escape from transmission of the ▶ genotype coding for it
predation, the higher the chance to transmit our will increase in frequency within a group of indi-
traits to the offspring. Any version of any herita- viduals. Consequently, it has been shown that
ble trait that favors a quick escape from predators cooperation between individuals increases with
should have been selected. Here we have the the genetic relatedness of the individuals. The
origin of mimicry. The more individuals resem- different cells composing a multicellular organ-
ble their substrate, the higher their chance of ism are genetically identical – we all are clones
survival against predation and thus the higher from the egg from which we were issued. Thus, in
the size of their expected progeny. Cryptic indi- sexually reproducing multicellular eukaryotes,
viduals (like the famous chameleon which can selection has produced a developmental system
adjust its color to the substrate’s one, or the in which most of our cells will not reproduce in
stick insects which are hardly distinguishable the long run because they die with the organism
from stems or leaves) transmit their heritable (the somatic cells constituting our body) while
traits that favor resemblance, which must have only a small part of the individual’s cells, the
been rapidly spread if the predation pressure sexual (or germinal) cells will survive through
was high. producing offspring. From an evolutionary per-
A fourth modality is the association of species. spective, the fact that one cell or another within
There is a gradient of situations between parasit- the organism produces a progeny makes no dif-
ism (association of species with a strong disequi- ference. Indeed, there can be competition among
librium in benefits) and mutualism (association of cells within an organism, but since only the ger-
species with mutual benefits). Our fleas are para- minal cells produce progeny, this process will not
sites. We have no benefit in hosting them. Our influence directly the hereditary make-up of the
mitochondria are mutualist (endo)symbionts: we next generation. It can, however, be important in
could not live without them inside our cells; they the determination of the success of the individ-
could not live without us. They are actually bac- uals in terms of survival or reproduction. For
teria whose ancestors got associated with eukary- instance, if some cells behave independently
otic cells more than 2,000 million years ago. from the others a cancer arises, with potentially
Almost all eukaryotes are symbiotic: they have immortal cell lineages but resulting in the death
mitochondria in their cells. ▶ Symbiosis is far of the organism.
more spread through life than we usually think. The same reasoning discussed in the above
All plants and photosynthetic algae are symbi- paragraph can easily explain parental care, since
otic: they host mitochondria and chloroplasts the hereditary traits of the parents will reproduce
(another organelle produced by bacterial endo- through the success of their progeny. It is inter-
symbiosis) in their cells. All vertebrate animals esting to note that in most species, the genetic
are symbiotic: we harbor a variety of bacterial relatedness between parents and offspring and
species in our digestive tract and we could not between siblings is equal. Consequently, in sev-
live without them. Symbiosis must be understood eral groups of organisms (termites, mole rats), the
individuals raise their sibs instead of producing When considering a structure in an organism,
their own progeny. In some species (insects we must be careful when evoking adaptation right
hymenoptera including bees, ants, wasps, etc.) away. Indeed organisms as wholes are compro-
however, the genetic relatedness is greater mises among traits. All traits are not under the
between sisters than between a mother and her same selective regime and “conflicts” occur. As
progeny. Hymenoptera are thus, not surprisingly, discussed above, there is no “optimal organism”
composed of a very high proportion of social to be understood as “perfect organism.” Perfec-
species where most individuals raise their sisters tion is not a scientific concept. Above all, the trait
and do not produce any progeny on their own. considered may not be present for direct selective
The family is thus a unit of selection as well as the reasons, but the by-product of a selection of
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individual. another trait of the same organism (see above).
Higher levels of selection also exist, but The side effect is obtained through chemical or
heated debates have been maintained concerning physical constraints of organism construction as
its application at the population level. One could consequences of a selection at another level or
think that populations that act in a coordinated trait. But there is another reason for adaptation
manner better face the changes of threats of the and side effects for a trait to be present: history.
environment than those that do not act A trait can be present as heritage or “frozen
so. Populations that protect their young can better accident” that has never been counter-selected.
transmit their traits to the offspring than those The pathway of our phrenic nerve is an example.
that do not. Populations that are more cohesive That nerve connects the diaphragm to our spinal
through help provided to the older or the weaker, cord. The pathway is not the shortest possible. It
through compassion, better face environmental passes through the thoracic cage, the neck and
changes. But theoretical models have shown connects the spinal cord at the level of the first
that, because individuals can migrate from one cervical vertebra. This long nerve is sometimes
population to another, this process can only act in irritated, provoking hiccup. That pathway is a
very particular cases. Indeed, imagine a popula- heritage of the ancient times, similar to the path-
tion of animals where everyone helps the others. way of the homologous nerve in teleosteans
If a migrant from a population where this trait (modern fish) that starts under the skull and inner-
does not exist arrives, he will benefit from the vates some muscular blocks of the branchial bas-
help without paying the cost of helping the others. ket situated just underneath. With the rise of
This issue is still debated. tetrapods (animals with limbs) some 360 Ma
Selection at the family level or at higher levels ago, the neck separated the muscular blocks
can be considered as providing an insight into the from the skull and throughout evolution the
natural origin of love, mutual help, compassion, length of the nerve increased. It is easy to under-
and moral rules, given that such behaviors are stand that the position where the nerve is getting
transmitted to the next generation, whatever the inserted in the spinal chord is more anatomically
nature of the transmission. Although it is too constrained than the progressive lengthening of
often forgotten, Darwin also wrote almost an the nerve as the muscular block it serves is getting
entire book (Darwin 1871) about this topic. The farther from the neck. The result is far from
ways by which our versions of our traits can be optimal but the interaction between selection
better spread through generations are diverse and and historical constraints have made it that way;
we should not reduce natural selection to a one of the effects of the above cited “evolutionary
“struggle for life” understood as a competition tinkering.”
among individuals. This probably does not reflect
the main factors by which natural selection plays, Evolution: Structures and History
and this surely does not reflect the richness of The processes described above explain why gen-
Darwin’s thinking and contemporary evolution- erations flow and change through time. That flow
ary thought. of generations is genealogy. A genealogy is a
network exhibiting ancestor-descendant relation- extinct, or expand by producing new branches.

ships between individuals. In the theoretical There is thus a selection at this level, but it can
scheme those relationships are among virtual only act by sorting out those lineages which do
individuals. In the empirical field those relation- possess characteristics making them sustainable.
ships are not accessible – except in the frame- These characteristics result from the processes
work of a few generations – because ancestors described above. Natural selection in its usual
have definitively disappeared. That flow of gen- sense is about the relative success of a given
erations is sometimes interrupted by extinctions, variation possessed by individuals, not about the
sometimes divided by obstacles. Obstacles can be relative success of species. Natural selection can
mountains (case of the domestic mice in the thus be seen as producing “locally optimal”
Madeira Islands), chromosomal changes, phero- organisms in terms of survival, or reproduction,
mone changes, behavioral changes, etc. For a or of transmission of their hereditary material, not
while, members of the genealogy of each part in terms of survival of the species.
cannot meet and give rise to offspring. If the Species definition should not be confused with
obstacle stays for long enough, alterations are criteria of assignment – both having been too
going to produce divergence. Divergence can be often confused into the single term of “species
strong enough to lead, sooner or later, to the concept.” When we find an individual organism
impossibility for the two sets to give rise again and we do not know what it is, we have criteria to
to offspring. The genealogy is a continuous flow assign the individual to an already known spe-
of changing individuals through times; at a small cies: similarity (most often), interbreeding, and
scale it is a network but at a larger scale there are fertile offspring. Note that sex, seen as gene
diverging branches (we will see later on that there exchange between two individuals, exists in all
can be connections among those branches under groups of organisms including Bacteria and
certain circumstances). Now we have to manage Archaea so that, even if the concept is less clear
another problem: we cannot speak about living in some groups, it is still possible to recognize
things by giving a surname to each individual we lineages which can, or cannot, mix their heredi-
meet. We need names at wider scales. We use tary material.
these branches to give names to sets of individ- Today we see sets of living beings that are
uals that are able together to give rise to off- separated by the impossibility of producing off-
spring. These sets are called species. The spring: in the history of biology species have
“concept” of species is more complex than it often been considered as real. The general theory
seems and an important literature has been pro- predicts that we must consider in the deep times
duced concerning it. Species has a definition: by that these sets are connected to each other: their
convention, a species is a set of individuals col- members do not produce offspring together but
lectively able to give rise to offspring, delimited they should have done it in the past. This will be
from one point of divergence in the genealogy to useful to reconstruct degrees of kinship among a
the next point of divergence. In other words, a given set of individuals. A cat has hair and a dog
species is an inter-node in the theoretical genea- has hair. Why do they have hair? Is it because
logical tree of life. A species seems to be real in they live here under this climate? No: the white
synchrony but is just conventional in diachrony bear and the giraffe have hair and they do not live
(i.e., considered through a significant time span): here. Do they have hair because they produce
today we know that cats do not make babies with offspring together? No: everyone knows that
dogs, but by considering the depth of times the dogs and cats, together, do not produce offspring.
theoretical genealogy does connect them and an But there has been a time when they did it. The
arbitrary limit must be set somewhere. Species is answer is in the past. Hair must have been
therefore defined as a convention of language inherited from common ancestors. By comparing
mapped onto a theoretical genealogical contin- organs, shape, metabolism, DNA sequences,
uum. Note that branches or the tree of life can go whatever feature that can be recognized as the
same in several individuals of different species, it together because they do share features in com-
can be interpreted as a sign of common ancestry. mon. The feature must define the group and must
By comparing many features, a tree can be not be present at the same time outside the group.
reconstructed, showing degrees in sharing fea- Because sharing features among species is
tures. Such a tree shows “who shares what with interpreted by the theory of evolution as common
whom.” It is not a genealogy because there are no ancestry, our classifications by shared features
ancestor-descendant relationships. When such a will be naturally extracted from phylogenies.
tree is read with the theory of evolution in mind, it A virtual “box” or “group” containing real organ-
becomes a ▶ phylogeny, a tree expressing “who isms is a taxon. By mapping our boxes onto nodes
is more closely related to who than others.” of a phylogeny, we produce complete taxa, i.e.,
E
A phylogeny does not express ancestor- taxa that contain one ancestor and all its known
descendant relationships between the actual indi- descendants. This is what we call monophyletic
viduals we have at hand, but expresses relative groups. In modern systematics – science of
degrees of relationships among them. There can classifications – classification of organisms is
be ancestors in a phylogeny, but they are virtual. related to the theory of evolution by the fact that
This is also true for fossils. Within a phylogeny, the only valid groups are monophyletic groups.
fossils of extinct species are treated like we treat The “tree of life” is a genealogical metaphor.
extant species. They are just a collection of fea- We will never have the full genealogy of life, just
tures gathered in a past organism. Fossils are at because ancestors have disappeared. But a theo-
the tip of branches just like extant organisms. retical scheme of genealogy is required by scien-
A given fossil can be the ancestor of some part tific evolutionary thinking. The shape of that
of today’s ▶ biodiversity, but we have no empir- theoretical tree is changing. Divergences of
ical mean to approach that kind of relationships. branches are best constrained for most of the
Modern phylogeny is not practiced anymore in animals; however for other eukaryotes, bacteria,
terms of “who is descending from whom” but in or archaea there can be exchange of material
terms of “who is closely related to whom,” which among well separated branches of the genealogy.
is more empirically powerful. Phylogenies are They are horizontal (also called lateral) transfers
used in a wide variety of scientific investigations. of genetic material. In bacteria it can be so pow-
Among them are classifications in natural history. erful that we are not sure that the shape of the tree
Classifications are conventional and classifica- is still the best scheme as a theoretical frame-
tion is the construction of a concept under the work. Another phenomenon we must be careful
form of a virtual box. Here outside there is no about is multiple symbioses. When partners of a
“bird.” Here outside there are only individuals. symbiosis are associated together for long
“Bird” is a name associated with a box in my enough, phylogenies reconstructed from each
mind, gathering millions of individuals, them- partner are expected to be the same. This is why
selves being gathered in some 10,000 species we use mitochondrial DNA to trace the phylog-
(themselves gathered in hundreds of genera, eny of eukaryotes. But when associations are
etc.). There are reasons why we created that recent, genetic traces of kinship can be different
box: a classification reflects properties of organ- for each of the partners.
isms we decided to point out. Did we decide to
put all these species in a same box because they
are good to eat? Because they are beautiful? Applications
Because they do the same things in their environ-
ment? Because they share the same features? Evolution Exported
A “good” or “efficient” classification is the one The discovery and understanding of evolution
that is self-consistent, i.e., that is successful in has had a number of consequences in the thought
organizing things according to the criteria we set. of our societies. It has changed the perception of
Today, systematists classify living organisms the world in many respects, influenced a number
of different scientific fields, and even been at the social and intellectual qualities or the “race” were
origin of new technologies. The fields where due to natural selection led the scientific commu-
evolution was exported are of two kinds. The nity to declare that there was a danger of degen-
first one is philosophical and sociological, i.e., eration due to the social laws adopted during the
outside scientific activity sensu stricto. It con- nineteenth century. The mix of political, socio-
cerns philosophy, eugenics, and social Darwin- logical, and scientific considerations led to the
ism. The second field is purely scientific: birth of an applied science: eugenics. It devel-
evolutionary thought was exported into already oped and was strongly supported by the scientific
developed scientific fields where it was initially community. In the USA and in Scandinavian
absent. This concerns evolutionary psychology, countries, thousands of people were sterilized in
Darwinian medicine, cellular Darwinism, bio- order to prevent their genes from spreading. The
chemistry, neural Darwinism, and genetic algo- Nazi government in Germany used this theory to
rithms in computer sciences. It would be too sterilize hundreds of thousands of people. These
long to develop these fields here but it is cer- practices disappeared more or less slowly after
tainly necessary to give some insights into some Nuremberg. Another social application was more
of them. economic, it was called “social Darwinism” in
From a philosophical point of view, once it is 1880 by Emile Gautier, a French journalist refer-
recognized that all living organisms are the result ring to the political philosophy of Herbert Spen-
of a natural process, the perception of Nature cer. It proposed the idea that the social system
(including human nature) is profoundly should respect the principle of selection and let
influenced. One of the major changes is that the the best succeed and the others remain desper-
fact that something exists does not anymore indi- ately poor in order to respect the laws of nature.
cate that it is good (because it is the will of the Note that, among the various modalities men-
Creator). The “Natural theology,” developed tioned above by which natural selection can
before Darwin (and which he was extensively play, Spencer considered only inter-individual
taught), loses its justification. As Thomas Huxley competition. This theory is still supported
(zoologist, friend of Darwin) stated, “Nature is among the theoreticians of neo-liberal economy.
neither moral nor immoral, but non-moral.” At Darwin himself never took a position in favor of
first sight, there are thus no moral rules to be Spencer’s ideas. Indeed, Darwin’s personal writ-
found in nature. However, since we are issued ings show that his political ideas were rather
from an evolutionary process, the sense of justice, different. On the left side of the political contin-
equity, and help could have something to do with uum, Kropotkine (1902) also referred to
the way the human lineage has evolved: there is a Darwin’s scientific writings when he tried to jus-
natural origin of our moral rules, a fact that is tify the naturalness of mutual help in human
perceived as contradicting their presumably tran- societies. It is important to point out that in either
scendental origin. Philosophical debates about case we have an ideological work at play, picking
these points have been going on for a century up into science what is interesting for socio-
and a half and we shall not enter them here. political purposes. These intellectual
Science is a collective activity whose first aim is by-products are not born by the sole will of the
just to rationally explain phenomena and objects scientific community trying to explain the natural
of the real world. As such it has no program to world.
contradict or confirm any theological or moral On a more positive side, in the second cate-
statement. gory of fields that have imported the idea of
From a social point of view, the Darwinian evolution, the modern theory of evolution can
theory, when fused with ▶ genetics under the help us understand psychology and the health
name “NeoDarwinism,” was first applied to the questions in humans; this has led to the develop-
human species. In the dark years following the ment of evolutionary medicine and psychology.
1929 economic crisis, the idea that the wonderful Moreover, the Darwinian concept of variation
selection has been imported into robotics, evo- just artificial selection among variants that
lutionary algorithms, protein engineering, lin- humans used to produce differential demographic
guistics, biochemistry, neurosciences, cellular success in their offspring.
biology, etc. Such fecundity leads to the ques- Evolution shapes medicine. A well-known
tion of whether, outside its initial field, the use of example is the rapid spread of antibiotic resis-
the word “evolution” should be restricted to the tance among different strains or species of bacte-
process of variation of entities and selection by ria by ▶ lateral gene transfer. It is produced by
their environment, or the word should still be the interchange of fragments of DNA (called
used in the wider vernacular sense of “a story plasmids) coding for proteins, able to degrade
to tell about change.” In the second sense there is an antibiotic used in the clinical setting. The
E
no doubt that the word “evolution” can be used plasmid that confers the antibiotic-resistant
to mean the change of our Universe and all ▶ phenotype in the recipient bacteria will be
objects it contains from galaxies to isolated mol- inherited by the progeny, allowing the resistant
ecules. In the first – more restrictive cells to evolve in the presence of such a selective
sense – applying the word evolution in the field pressure. A second example: when people
of astronomy implies the identification of extra- infected by the human immunodeficiency
terrestrial objects susceptible to variation and ▶ virus (HIV) are treated with antiviral drugs,
the transfer of structure to similar entities, a the population of viruses that infects them
transfer subject to environmental conditions. evolves multiple amino acid substitutions that
Thus, this process has not been used to describe confer drug resistance in few months. As a con-
the change of most of the large-scale objects we sequence of HIV evolution, the antiviral drug
find in the Universe. However the word “evolu- combination must be changed.
tion” has been used to describe changes in mac- Usually, evolution is not directly accessible to
romolecules that could be present in meteorites our senses, since large animals have long gener-
or planets, like amino acids or nucleotide poly- ation times and require very long periods to
mers. Biological macromolecules were, what- exhibit spectacular changes or speciation events.
ever the ways by which they transfer their In turn, organisms with very short generation
variations in sequence or conformation to others, times, such as Escherichia coli and other bacte-
probably the very first step of (precellular) ria, evolve very fast but we cannot see them with
evolution in the restrictive sense. As at least our sole eyes. Nevertheless, the evolution of fast-
small precursors of the macromolecules are growing viruses or bacteria is easily monitored in
present in our extraterrestrial Universe, “evolu- the lab. Additionally, thanks to the advances in
tion” is a rightful and suggestive concept in molecular biology, in vitro evolution is currently
astrobiology. used to mimic and speed up the evolutionary
processes at the molecular level.
Evolution Everyday Last but not least, evolution does not mean nor
Evolution is not only a matter of fossils or DNA justify anything in terms of political or moral
sequences. Evolution is needed to feed the entire issues. Evolution is just the scientific approach
human kind. There would be no agriculture with- to understand the origin of living beings and their
out evolution. Artificial selection helped us to changes through time. The approach uses amoral
survive by domesticating various animals and procedures, and our moral considerations do not
plants. Often, one species has yielded very differ- have to be transferred to the living world (and
ent varieties. The sole species Brassica oleracea reciprocally). Each time that such a transfer
yielded cabbage, broccoli, cauliflower, Brussels occurred along the history of our evolutionary
sprouts, and flowering kale. The different breeds sciences, it was a source of obstacles. Moral and
of dogs (the species Canis lupus familiaris) differ political issues must be managed in the philo-
much more in morphology and behavior than do sophical and political realms that surround scien-
other mammalian species. This is also evolution: tific activity. They can give directions to
780 Evolution, Chemical
research; however, scientific reasoning itself

must stay out of these constraints. Evolution, Chemical
André Brack
See Also Centre de Biophysique Moléculaire CNRS,
Orléans Cedex 2, France
▶ Adaptation
▶ Biodiversity
▶ Cell Keywords
▶ Earth’s Atmosphere, History of the Origins Autotrophic life; Cellular world; Primitive life;
▶ Evolution, In Vitro Primordial soup; RNA world
▶ Fitness
▶ Genetics
▶ Genotype Synonyms
▶ Lateral Gene Transfer
▶ Life Prebiotic chemistry
▶ Multicellular Organisms
▶ Mutation
▶ Natural Selection Definition
▶ Phenotype
▶ Phylogenetic Tree Chemical evolution refers to the suite of natural
▶ Phylogeny reactions that led to the first living systems from
▶ Selection abiotically synthesized molecules on the primi-
▶ Species tive Earth. Since this was a historical process and
▶ Symbiosis we have no relics of the compounds formed in
▶ Taxonomy this process, chemists can only model chemical
▶ Virus evolution by running experiments, chemical
reconstructions also known as prebiotic
References and Further Reading chemistry.
Barton NH, Briggs DEG, Eisen JA, Goldstein DB, Patel

NH (2007) Evolution. CSHL Press, Cold Spring Har- Overview
bor, p 833
Darwin C (1859) On the origin of species. John Murray,
London Although primitive life is generally believed to
Darwin Ch (1871) The descent of man, and selection in have been organic, that is, based on ▶ carbon
relation to sex, 2 vols. John Murray, London chemistry, the precise sequence of steps that
Kropotkine P (1902) Mutual aid: a factor of evolution.
Porter Sargent Handbooks, London
allowed for the formation of living systems
Lecointre G, Le Guyader H (2006) The tree of life, a from abiotic chemistry remains poorly
phylogenetic classification. Harvard University Press, constrained and somewhat speculative. Chemi-
Cambridge, p 560 cal evolution is the study of the processes that
Lyell C (1832) Principles of geology, vol 2. John Murray,
London
led from simple molecules to biochemistry.
Milner R (2009) Darwin’s universe: evolution from A to There are presently numerous scenarios for
Z. University of California Press, Los Angeles, p 488 this process which scientists consider plausible.
Spencer H (1862) A system of synthetic philosophy, Among them is the idea of a genetic mineral
vol I. Williams & Norgate, London
Tort P (dir) (1996) Dictionnaire du Darwinisme et de
material presenting suitable properties such as
l'évolution, 3 vols. Presses Universitaires de the ability to store and replicate information
France, Paris (Cairns-Smith 1982). Another hypothesis favors
Evolution, In Vitro 781
a heterotrophic origin of life in a “▶ primordial Morowitz H (1992) Beginnings of cellular life. Yale Uni-
soup,” involving self-assembling, preformed versity Press, New Haven
Wächtershäuser G (2007) On the chemistry and evolution
carbon-based molecules. An autotrophic origin of the pioneer organism. Chem Biodiv 4:584–602
of life via direct reduction of carbon oxides in a
“metabolism-first” scenario has also been pro-
posed (Wächtershäuser 2007).
While it is clear that all contemporary life
shares a common heritage, it is possible that Evolution, In Vitro
there were earlier, more primitive states that
life evolved through which might have borne Carlos Briones E
little resemblance to modern biochemistry. Centro de Astrobiologı́a (CSIC/INTA), Consejo
Some believe that primitive life emerged as a Superior de Investigaciones Cientı́ficas, Madrid,
cell-like system requiring at least pre-RNA mol- Spain
ecules capable of storing and transferring the
information needed for reproduction,
pre-enzymes providing the basic chemical Keywords
work, and pre-membranes able to isolate the
system from the aqueous environment. Since Amplification; Aptamer; Combinatorial nucleic
RNA has been shown to be able to act as an acid library; Deoxyribozyme; Evolution; Molec-
information molecule and also as a catalytic ular evolution; Mutation; Ribozyme; RNA
molecule, RNA has been considered as a candi- world; Selection; SELEX ®
date for the first living system that preceded the
cellular world. The spontaneous organization of
amphiphilic molecules to form vesicles has also
been postulated as the first step toward the origin
Synonyms
of life (Morowitz 1992). Chemists are also
Artificial evolution; Directed evolution; Experi-
tempted to consider that primitive self-
mental evolution; In vitro evolution
replicating systems depended on simple autocat-
alytic molecules adsorbed on solid surfaces,
which could solve some of the problems of the
likely high dilution of organics in the primitive Definition
oceans.
In vitro evolution is an experimental method that
allows the screening of large random-sequence
See Also libraries of nucleic acids – ▶ RNA or
▶ DNA – for a specific function, such as cataly-
▶ Abiotic sis or binding. In vitro evolution experiments
▶ Carbon have led to the selection of novel or improved
▶ Earth’s Atmosphere, History of the Origins RNA enzymes, called ▶ ribozymes, as well as
▶ Origin of Life artificial DNA enzymes – DNAzymes or
▶ Prebiotic Chemistry deoxyribozymes – and target-binding nucleic
▶ Primordial Soup acid molecules, ▶ aptamers. The iterative
▶ RNA World in vitro process involves serial cycles of ampli-
fication and selection of the functional nucleic
acid until the desired activity is reached. The
outcome of directed evolution can be monitored
Cairns-Smith AG (1982) Genetic takeover. Cambridge through the analysis of the genotype and pheno-
University Press, Cambridge type of the evolved molecules.
782 Evolution, In Vitro
History directed evolution of ribozymes (Wright and

Joyce 1997). In parallel, a new method termed
The first experiments aimed at determining if “mRNA display” was set up for the in vitro selec-
Darwinian evolution was possible in cell-free tion of peptides and proteins (Roberts and
systems involved the in vitro replication of the Szostak 1997).
RNA genome of the bacteriophage
Qb – a ▶ virus that infects Escherichia coli – by
the viral RNA replicase protein. Serial dilution Overview
experiments allowed the transfer of an aliquot of
the replication mixture into a new reaction tube The starting point of a stepwise in vitro evolution
containing fresh enzyme and RNA monomers. experiment is a large pool of nucleic acid mole-
The inherent mutation rate of the viral polymer- cules, in general, 1012–1016, with a central region
ase ensured genetic variation during the experi- of random or mutagenized sequence, usually
ment, and the ▶ phenotype under selective 20–100 nt long, flanked by two primer-binding
pressure was the replication speed of the RNA sites of defined sequence. The successive
genome. After several serial transfers, the Qb amplification-selection rounds increase the ratio
genome evolved by deleting sequences not of active:inactive RNA – or DNA – and the pool
required for its recognition by the polymerase, finally becomes dominated by molecules with the
thus shortening the replication time (Mills et al. desired ▶ phenotype. Within each round, RNA
1967). amplification involves successive steps of reverse
A quarter century later, the availability of transcription, PCR amplification, and ▶ tran-
reverse transcriptase enzyme – RT, which cata- scription. ▶ Selection is introduced by choosing
lyzes the conversion of RNA into DNA – and for the next round only the nucleic acid molecules
PCR technology, as well as the possibility to able to bind the target – aptamers – or to perform
chemically synthesize nucleic acid pools of ran- the desired activity – ribozymes – while
dom sequence, opened a new era in in vitro evo- nonfunctional molecules within the pool are
lution experiments. This field was also reinforced discarded. The whole cell-free process occurs
by the discovery of natural ▶ ribozymes, what in vitro, and all the experimental variables can
underlined the functional capabilities of RNA be controlled and fine-tuned. In vitro evolution
and its hypothesized key role in pre-cellular evo- can be distinguished from in vitro selection in
lution. By 1990, directed evolution allowed that the former includes a continuous introduc-
the – artificial – selection of functional RNA tion of genetic variation in the molecular popula-
molecules from random pools in two independent tion, through mutation – usually, by error-prone
experiments that used the ability of RNA to bind PCR – and/or recombination (Wilson and
a target molecule as the selective pressure. The Szostak 1999; Joyce 2004).
stepwise process of amplification-selection was In vitro evolution and in vitro selection have
termed “Systematic Evolution of Ligands by been used to obtain artificial nucleic acid
Exponential Enrichment” or SELEX (Tuerk and enzymes and to engineer natural ribozymes to
Gold 1990), and the term “aptamer” was coined expand or improve their current functionality.
to denote the in vitro evolved, target-binding The unveiled functional plasticity of RNA has
RNA (Ellington and Szostak 1990). increased the plausibility of the ▶ RNA World
The next milestone in experimental cell-free hypothesis for the origin and early evolution of
evolution was the development of a system for life (Joyce and Orgel 2006). In parallel, directed
continuous in vitro evolution of nucleic acids. In evolution of nucleic acids has generated new
contrast to stepwise in vitro evolution, it com- families of aptamers with diagnostic and thera-
bined the processes of selection, amplification, peutic applications (Klussmann 2006).
and genetic diversification within the same reac- Microfluidic chip technology and computer-
tion mixture, thus allowing the continuous assisted control have been incorporated to the
Evolution, Molecular 783
biochemical process in order to automate the

system and to monitor in real time the population Evolution, Molecular
size, its composition, and the performance of the
evolved molecules (Paegel and Joyce 2008). Peter Schuster
Institut f€ur Theoretische Chemie der Universität
Wien, Wien, Austria
See Also
▶ Aptamer Keywords
▶ DNA E
▶ Evolution, Biological Adaptive evolution; Applied molecular evolu-
▶ Evolution, Molecular tion; Error threshold; Evolutionary biotechnol-
▶ Genotype ogy; Genome; Genotype; Neutral evolution;
▶ Phenotype Phenotype; Phylogenetics; Quasispecies;
▶ Polymerase Chain Reaction SELEX ®
▶ Quasispecies
▶ Ribozyme
▶ RNA Synonyms
▶ RNA World
▶ Selection Directed evolution; Molecular phylogenetics
▶ Transcription
▶ Virus
Definition
References and Further Reading Molecular evolution deals with the application of
Ellington AD, Szostak JW (1990) In vitro selection of
knowledge from biophysics, biochemistry, and
RNA molecules that bind specific ligands. Nature molecular biology to biological evolution with
346:818–822 the focus on understanding evolutionary phe-
Joyce GF (2004) Directed evolution of nucleic acid nomena at the molecular level. Two branches of
enzymes. Annu Rev Biochem 73:791–836
Joyce GF, Orgel LE (2006) Progress toward understand-
molecular evolution are established fields of
ing the origin of the RNA world. In: Gesteland RF, research: (1) reconstruction of phylogenetic
Cech TR, Atkins JF (eds) The RNA world. Cold trees by comparing genomic DNA sequences
Spring Harbor Laboratory Press, New York and (2) modeling evolution in vitro and in silico
Klussmann S (ed) (2006) The aptamer handbook. Wiley-
VCH, Weinheim
(i.e., by computer simulation), using simple
Mills DR, Peterson RL, Spiegelman S (1967) An extra- models like populations of RNA molecules, sim-
cellular Darwinian experiment with a self-duplicating ple viruses, or bacteria.
nucleic acid molecule. Proc Natl Acad Sci U S A
58:217–224
Paegel BM, Joyce GF (2008) Darwinian evolution on a
chip. PLoS Biol 6, e85 History
Roberts RW, Szostak JW (1997) RNA-peptide fusions for
the in vitro selection of peptides and proteins. Proc Molecular evolution originated from the vision of
Natl Acad Sci U S A 94:12297–12302
an evolutionary molecular clock by Linus Pau-
ligands by exponential enrichment: RNA ligands bo ling and Emile Zuckerkandl in 1962 (Morgan
bacteriophage T4 DNA polymerase. Science 249: 1998). Reconstruction of phylogenetic or evolu-
505–510 tionary trees by comparison of biomolecular
Wilson DS, Szostak JW (1999) In vitro selection of func-
tional nucleic acids. Annu Rev Biochem 68:611–647
sequences became an established method through
Wright MC, Joyce GF (1997) Continuous in vitro evolu- the works of Walter Fitch, Emanuel Margoliash,
tion of catalytic function. Science 276:614–617 and Margaret Dayhoff in the 1960s, and the
784 Evolution, Molecular
evolutionary clock (Kumar 2005) became an the tree of life from present-day sequences of
established tool in evolution research for dating species (Graur and Li 2000) in which the proper
speciation events. In vitro evolution of RNA alignment of sequences is an important part of
molecules from bacteriophages was initiated by the procedure (Mount 2004). Comparing
Sol Spiegelman in the 1960s (Spiegelman 1971). sequences from vertebrates reveals the molecu-
He showed that Darwinian evolution occurs in lar clock of evolution (Morgan 1998; Kumar
cell-free systems. At the same time, Manfred 2005; Takahata 2007), which implies that the
Eigen developed a kinetic theory of evolution at number of single point mutations per nucleotide
the molecular level (Eigen 1971). Since 20 years, site and time is approximately constant for a
molecular evolution has been applied to the evo- given protein molecule. Further investigations
lutionary design of molecules with predefined have shown substantial vagaries of the clock,
properties and given rise to a new branch of but in the age of genomics, the molecular clock
biotechnology. turned out to be a valuable device for dating
historical events. The last 40 years have seen a
heavy debate between selectionists and neutral-
Overview ists about the role of mutations that have no
influence on the ▶ fitness values of their carriers.
Evolution is based on the interplay of reproduc- Data on the evolution of entire genomes are
tion (providing the basis for inheritance, either by ending this controversy through the discovery
variation or ▶ mutation as a result of imperfect of new mechanisms dominating genome evolu-
copying or recombination of genetic modules) tion (Nei 2005).
and selection (choosing from the phenotypes pre- In the language of chemical kinetics, the evo-
sent in the population). Insights into the molecu- lution of molecules through correct replication
lar machinery of life reveal the specific chemistry and mutation under controlled conditions
of reproduction initiated by replication of a (Fig. 1) is cast into a first-order differential
nucleic acid molecule – DNA, or rarely equation:
RNA – which carries the genetic information in
the form of a one-dimensional string of digits. dxj Xn
Mutation occurs in three different forms: (1) sin- ¼ Q f x xj ’ðtÞ with
i¼1 ji i i
dt Xn Xn (1)
gle nucleotide incorporation errors commonly ’ðtÞ ¼ f x and x ¼ 1:
i¼1 i i i¼1 i
called point mutations, (2) insertions, when a
part of the sequence is copied twice or several
This equation describes the changes in geno-
times, and (3) deletions, when a part of the
type distribution of a population that consists of
sequence is missing in the copy. The unfolding
n different genotypes X1, X2, . . ., Xn. The vari-
of the genetic information, the ▶ genotype, or the
ables are relative concentrations:
genome, yielding the ▶ phenotype, is the key to Xn
understanding evolution, because it creates the xj ¼ N j = i¼1 N i with Ni = [Xi] being the num-
repertoire upon which selection operates. The ber of molecules of type Xi. The parameters in
genotype-phenotype map is the source of com- Eq. 1 are (1) the replication rate parameters or
plexity and varies from sequence-structure rela- fitness values for the individual genotypes, fi, and
tions for in vitro evolution of molecules to the (2) the mutation frequencies, Qji representing the
entire process of development leading from the frequency at which Xj is obtained as an error copy
fertilized egg to the adult organism in case of of the template Xi. The first term in Eq. 1
higher organisms. describes the production of Xj either as a correct
Every organism carries information on its copy, Qjjfjxj, Xor as an error copy from another
n
own history in the form of the DNA or RNA genotype Xi, Q f x . The second term,
i¼1, i6¼j ji i i
sequence that originates through mutation ’(t), is a flux compensating for the excess pro-
events. Molecular phylogenetics reconstructs duction of genotypes. Mathematical analysis of
Evolution, Molecular, Fig. 1 An example of a reaction inheritance requires Qjj > Qij and sufficiently large (see
mechanism for replication and mutation according to the error-threshold shown in Fig. 2). Since every replica-
Eq. 1. An enzyme binds the template molecule (Xj) and tion product is either correct or a mutant, a conservation
the building blocks for polynucleotide synthesis (M). Cor- relation Si Qij = 1 holds. After the replication is com-
rect replication and mutation are parallel chemical reac- pleted both polynucleotide molecules, template and copy
tions with a rate parameter fj, individual reaction channels are released from the enzyme, which is free for the next
lead to correct replication with frequency Qjj and to a replication round
variety of mutants with frequency Qij with i 6¼ j. Stable
the solutions of Eq. 1, which are obtained by mutational backflow from mutants to the mas-
integrating factor transformation and solution of ter sequence, the critical mutation rate is
an n n eigenvalue problem (Eigen et al. 1989), obtained as
yields two results that are relevant for biology:
pcr ¼ 1 s1=‘
m with
1. The long time or stationary solution is given
fm
by the largest eigenvector x0 of the matrix sm ¼ and
fm (2)
W = QF, where Q = {Qij; i, j = 1,. . ., n}
and F = {Fij = fi dij; i, j = 1,. . ., n}, the diag- 1 Xn
fm ¼ fx :
i¼1, i6¼m i i
onal matrix of fitness values (dij is 1 xm
Kronecker’s symbol, dij = 1 if and only if
i = j and 0 otherwise). All elements of x0 are In this equation, the chain length of the geno-
positive by Perron-Frobenius theorem and type is denoted by ‘, xm is the relative concentra-
hence no genotype will vanish. The largest tion of the master sequence, sm is the superiority
eigenvalue l0 is nondegenerate and hence the of the master sequence, and fm is the mean fitness
result of the mutation selection process is of the population except the master sequence.
unique. No matter what the initial conditions Equation 2 has two important consequences: It
were, the population will converge toward a defines (1) a maximal chain length for constant
unique distribution of genotypes called replication accuracy (1 p), ‘max
lnsm =p, and
quasispecies (Eigen and Schuster 1979), (2) a maximal error rate for constant chain length
which consists of a fittest genotype, the master ‘, pmax
lnsm =‘. Both limitations have immedi-
sequence Xm, and its mutant cloud. ate consequences in biology: (1) In classes of
2. Consideration of quasispecies as a function of organisms, the spontaneous mutation rates deter-
the mutation rate p, x0 (p), reveals a critical mined by the accuracy of the replication mecha-
mutation rate pcr at which the genotype distri- nism are inversely proportional to genome
bution changes abruptly from the quasispecies lengths (Drake et al. 1998; Gago et al. 2009),
centered around the master sequence into the and (2) for RNA virus replication, which occurs
uniform distribution (Fig. 2). Neglecting at mutation rates as close to the error threshold as
786 Evolution, Molecular
Relative concentration y(p)

0.8
0.6
0.4
0.2
0 0.01 0.02 0.03 0.04 0.05

Mutation rate p
0.08
Relative concentration y(p)
0.06
0.04
0.02
0.016 0.018 0.02 0.022 0.024 0.026 0.028 0.03
Mutation rate p
Evolution, Molecular, Fig. 2 The accuracy limit for the sum of the concentrations of all three-error mutants,
replication. The two plots show solution curves of Eq. 1 etc., and in general yk(p) is the sum of the concentrations
as functions of the mutation rate p. In order to illustrate the of all k-error mutants. At the error-threshold the
phase transition-like change in the mutant distribution at quasispecies changes abruptly into the uniform distribu-
the error threshold (2) the relative concentration of error tion, and the concentrations of the mutant classes become
classes yk(p) are plotted against p: y0(p) represents the equal to the binomial coefficients. The lower part of the
concentration of the master sequence, y1(p) is the sum of figure represents an enlargement of the curves around the
the concentrations of all one-error mutants, y2(p) is the error-threshold. Parameters used in the calculations: chain
sum of the concentrations of all two-error mutants, y3(p) is length ‘ = 100, fitness values f0 = 10, fn = 1 for all n 6¼ 0
possible in order to guarantee maximal variabil- Evolution experiments under controlled condi-
ity in fighting the defense system of the host, a tions were also carried out with bacteria, and
drug-induced increase of the mutation rate can radical innovations induced by mutations were
drive the population into extinction (lethal muta- interpreted successfully at the molecule level
genesis; Domingo et al. 2008). (Blount et al. 2008). The molecular approach to
Molecular evolution experiments starting with evolution is completed by introducing a relation
the seminal work of Sol Spiegelman (Spiegelman between genotypes and phenotypes in the form of
1971) and continued by Christof Biebricher, Jack a mapping or fitness landscape, which in the
Szostak, Gerald Joyce, and others (Watts and simplest system, in vitro evolution of RNA,
Schwarz 1997; Joyce 2004, 2007) represent deals with RNA sequences and structures
experimental realizations of the theory. (Schuster 2003, 2006). Evolutionary phenomena,
for example, the existence of error thresholds, are Applications

highly sensitive to the fitness landscape and com-
monly applied. Oversimplified models based on In vitro evolution of molecules was applied to the
the unrealistic assumptions that mutations con- design of molecules with predefined properties in
tribute to fitness additively or in a multiplicative biotechnology and became a popular tool for
way give wrong results (Phillipson and Schuster systematic evolution of ligands by exponential
2009). Neutrality is a fundamental property of enrichment (SELEX ®, which is a registered
realistic fitness landscape, and neutral evolution trademark of the company Gilead Sciences, Fos-
(Kimura 1983) in parts of the sequence space is ter City, CA). Examples are the evolutionary
indispensable for the success of the Darwinian design of nucleic acid molecules (Joyce 2004;
E
mechanism (Schuster 2006). Klussmann 2006), proteins (Brakmann and
Molecular kinetics of evolution and most of Johnsson 2002; Jäckel et al. 2008), and synthetic
population genetics are based on modeling by compounds (Wrenn and Harbury 2007).
means of differential equations. The limitations Quasispecies theory and the concept of error
of this approach primarily come from small par- propagation in successive replication gave rise
ticle numbers – every mutant has to start from a to new antiviral strategies known as lethal muta-
single copy; uniform distributions are completely genesis (Domingo et al. 2008). Together with
unrealistic since there are orders of magnitude rational design, evolutionary biotechnology rep-
less molecules in the population than possible resents a core technique of synthetic biology.
genotypes. Proper stochastic treatments and com-
puter simulations were performed, but a universal
model of finite population size effects in evolu- Future Directions
tion is not at hand.
The promising and challenging future program
for molecular evolution is the integration of data
Basic Methodology from genomics and systems biology into the cur-
rently used methods. Whereas phylogenetics on
The methodology of molecular evolution is the whole genome level is more or less solved,
shared with molecular genetics, genome upscaling of modeling evolution from RNA mol-
research, population genetics, bioinformatics, ecules and viruses to cells and organisms will
and biochemical kinetics. Specific developments require new experimental and computational
concern the construction of special flow reactors tools. Learning how genotypes are mapped into
and other equipment for evolutionary biotechnol- phenotypes represents the key for understanding
ogy (Watts and Schwarz 1997). evolution at the molecular level.
Key Research Findings See Also
The phylogenetic tree of conventional biology ▶ Evolution, Biological

has been approved independently by molecular ▶ Evolution, In Vitro
phylogenetics. The molecular clock of evolution ▶ Fitness
yields a genetic measure of time. In vitro evolu- ▶ Flow Reactor
tion has provided evidence that Darwinian evo- ▶ Genotype
lution does not require reproduction at the ▶ Hypercycle
cellular level. Quasispecies theory and existence ▶ Mutagenesis
of error thresholds for replication provide a solid ▶ Mutation
theoretical basis for virus evolution at the molec- ▶ Phenotype
ular level. ▶ Phylogenetic Tree
788 Evolutionary Algorithms
▶ Phylogeny Evolutionary dynamics. Exploring the interface of

▶ Quasispecies selection accident, neutrality, and function. Oxford
University Press, Oxford, UK, pp 163–215
▶ Replication (Genetics) Schuster P (2006) Prediction of RNA secondary struc-
tures: from theory to models and real molecules. Rep
Prog Phys 69:1419–1477
Spiegelman S (1971) An approach to the experimental
References and Further Reading analysis of precellular evolution. Q Rev Biophys
4:213–253
Blount ZD, Borland CZ, Lenski RE (2008) Innovation by Takahata N (2007) Molecular clock: an anti-neo-
mutation in long time evolution of Escherichia coli in Darwinian legacy. Genetics 176:1–6
constant environment. Proc Natl Acad Sci U S A Watts A, Schwarz G (eds) (1997) Evolutionary
105:7899–7906 biotechnology – from theory to experiment. Biophys
Brakmann S, Johnsson K (eds) (2002) Directed molecular Chem 66:67–290
evolution of proteins. How to improve enzymes for Wrenn SJ, Harbury PB (2007) Chemical evolution as a
biocatalysis. Wiley-VCh, Weinheim tool for molecular discovery. Annu Rev Biochem
Domingo E, Parrish CR, Holland JJ (2008) Origin and 76:331–349
evolution of viruses, 2nd edn. Academic/Elsevier,
Amsterdam
Drake JW, Charlesworth B, Charlesworth D, Crow JF (1998)
Rates of spontaneous mutation. Genetics 148:1667–1686
Eigen M (1971) Self-organization of matter and the evo-
lution of biological macromolecules. Naturwis- Evolutionary Algorithms
senschaften 58:465–523
Eigen M, Schuster P (1979) The hypercycle. A principle ▶ Genetic Algorithms
of natural self-organization. Springer, Berlin
Eigen M, McCaskill J, Schuster P (1989) The molecular
quasispecies. Adv Chem Phys 75:159–264
Gago S, Elena SF, Flores R, Sanjuán R (2009) Extremely
high mutation rate of a hammerhead viroid. Science
323:1308
Evolutionary Sequence of Young
Graur D, Li WH (2000) Fundamentals of molecular evo- Stellar Objects
lution, and Li WH (2006) molecular evolution. Sinauer
Associates, Sunderland ▶ Spectral Classification of Embedded Stars
Jäckel C, Kast P, Hilvert D (2008) Protein design by
directed evolution. Annu Rev Biophys 37:153–173
Joyce GF (2004) Directed evolution of nucleic acid
enzymes. Annu Rev Biochem 73:791–863
Joyce GF (2007) Forty years of in vitro evolution. Angew
Chem Int Ed 46:6420–6436 Evolutionary Tree
Kimura M (1983) The neutral theory of molecular evolu-
tion. Cambridge University Press, Cambridge, UK ▶ Phylogenetic Tree
Klussmann S (ed) (2006) The aptamer handbook. Func-
tional oligonucleotides and their applications. Wiley-
VCh, Weinheim
Kumar S (2005) Molecular clocks: 4 decades of evolution.
Nat Rev Genet 6:654–662
Morgan GJ (1998) Emile Zuckerkandl, Linus Pauling, and
Exergonic
the molecular evolutionary clock, 1959–1965. J Hist
Biol 31:155–178 Jacques Reisse
Mount DW (2004) Bioinformatics, 2nd edn, Sequence and Université Libre de Bruxelles, Brussels, Belgium
genome analysis. Cold Spring Harbor Laboratory
Press, Cold Spring Harbor
Nei M (2005) Selectionism and neutralism in molecular
evolution. Mol Biol Evol 22:2318–2342 Definition
Phillipson PE, Schuster P (2009) Modeling by nonlinear
differential equations. Dissipative and conservative
processes. World Scientific, Singapore, pp 9–60
The term exergonic refers to a chemical reaction
Schuster P (2003) Molecular insights into evolution of that releases free energy as it proceeds. An exer-
phenotypes. In: Crutchfield JP, Schuster P (eds) gonic reaction may also be ▶ exothermic, but
Exobiologie Experiment 789
does not need to be (see ▶ entropy). An example

of exergonic reaction is gasoline ▶ oxidation in
the Earth’s atmosphere; burning gasoline releases
sufficient free energy (work) to propel a car.
It is important to make a clear distinction
between thermodynamics and kinetics.
A spontaneous reaction can be characterized by
a rate so small that no change can be observed, as
is the case with gasoline in the presence of oxy-
gen at normal temperature. A temperature
E
increase by a flame or spark is sufficient to initiate
a combustion reaction which, being not only
exergonic but also exothermic, then proceeds
without any additional supply of heat. Many bio-
chemical reactions are exergonic but extremely
slow in the absence of a catalyst (for example an
enzyme). In the presence of enzymes, however,
the reactions proceed at an appreciable rate at low
temperature.
Exobiologie Experiment, Fig. 1 Experiment

“exobiologie” as seen from the interior of the Kvant mod-
See Also ule of the MIR space station (Photo CNES)
▶ Entropy
deposited on windows of magnesium fluoride and
▶ Exothermic
exposed directly to the solar UV flux. Spores of
▶ Oxidation
Bacillus subtilis mixed with meteoritic powder
were also exposed in cooperation with the
DLR. As references, duplicates of the samples
were placed in the instrument but hidden from the
Exobiologie Experiment solar light as ground-based experiment was
conducted.
Michel Viso Analysis of the material after the flight did not
CNES/DSP/SME, Vétérinaire/DVM, reveal any racemization or polymerization of the
Astro/Exobiology, Paris Cedex 1, France amino acids but did provide information regard-
ing photochemical pathways for the degradation
of leucine and of the tripeptide. Very thin layers
Definition of inorganic material did not protect spores
against the deleterious effects of energy-rich
During the ▶ CNES sponsored mission “Per- UV radiation in space to the expected amount.
seus,” the experiment “exobiologie” has been Surprisingly, self-formed layers of UV radiation
flown in 1999 onboard the Russian MIR space inactivated spores serve as a UV shield by them-
station and exposed for 97 days outside the sta- selves. The hypothetical interplanetary transfer
tion (April 16–July 9). Samples containing chi- of life by the transport of microorganisms inside
ral ▶ amino acid (▶ leucine and alpha-methyl rocks through the solar system cannot be
leucine) and peptides (leucine-diketopiperazine excluded but requires the shielding of a substan-
and trileucine thioethylester), mixed or not with tial mass of inorganic substances (Rettberg
montmorillonite or meteoritic powder, were et al. 2002; Fig. 1).
790 Exobiology

Exogeny
Boillot F, Chabin A, Buré C, Venet M, Belsky A,
Jacquet R, Bertrand M, Delmas A, Brack A, Barbier
▶ Exogenous
B (2002) The PERSEUS exobiology mission on MI-
R. Behaviour of amino acids and peptides in Earth
orbit. Orig Life Evol Biosph 32(4):359–385
Rettberg P, Eschweiler U, Strauch K, Reitz G, Horneck G,
Wänke H, Brack A, Barbier B (2002) Survival of
microorganisms in space protected by meteorite mate-
rial: results of the experiment ‘EXOBIOLOGIE’ of the ExoMars
PERSEUS mission. Adv Space Res 30(6):1539–1545
Jorge L. Vago
European Space Agency – ESA/ESTEC
(SRE-SM), Noordwijk, The Netherlands
Exobiology
▶ Astrobiology Keywords
ESA; Habitability; Life on Mars; Mars evolution;

Mars Express; Mars geology; Mars missions;
Exogenous Mars rovers; MER; MSL; Methane on Mars;
MRO; Roscosmos; NASA Pathfinder; Viking;
Daniele L. Pinti Water on Mars
Montréal, QC, Canada Definition
ExoMars is a cooperative program of the

Synonyms ▶ European Space Agency (ESA) and the Rus-
sian Federal Space Agency, Roscosmos. Two
Exogeny missions are foreseen within the ExoMars pro-
gram for the 2016 and 2018 launch opportuni-
ties to ▶ Mars. The 2016 ExoMars Trace Gas
Definition Orbiter (TGO) mission will study Martian
atmospheric trace gases, such as methane, to
In geology, exogenous refers to all the processes acquire information on possible active geologi-
that are produced at the surface of the Earth (and cal or biological processes. The observations
other planets). ▶ Weathering, erosion, transport, will target sources, distribution, and temporal
and sedimentation are the main exogenous variation of various atmospheric species. This
processes. The result of these processes is the mission will also deliver an Entry, Descent, and
formation of sediments and ▶ sedimentary rocks. Landing Demonstrator Module (EDM) to the
surface of Mars. The 2018 mission will deliver
the ExoMars rover to the surface of Mars.
See Also ESA’s ExoMars rover will be devoted to
the search for signs of past and present life,
▶ Endogenous using a subsurface drill and a novel payload.
▶ Sedimentary Rock The lander will be equipped with additional
▶ Weathering instruments.
ExoMars 791
History to develop and launch two ExoMars missions

in 2016 and 2018.
ExoMars was originally conceived as a rover The 2016 mission consists of two major
mission carrying a drill capable to collect elements: (1) the Trace Gas Orbiter (TGO)
small samples from the subsurface and analyze that will search for evidence of methane and
them. The initial work aiming to define the other atmospheric gases to acquire information
scope of the rover mission and its payload on possible active geological or biological pro-
started in 1996, when a group of scientists was cesses (the TGO will also serve as a data relay
tasked by ESA to make recommendation for for surface missions until end 2022) and (2) the
future search-for-life missions on Mars and the Entry, Descent, and Landing Demonstrator
E
rest of the Solar System. Their findings were Module (EDM) that will land on Mars to vali-
collected in the so-called ESA “Red Book” date key technologies for the 2018 mission.
report of 1999. During 2001, the nascent ESA The 2018 mission will deliver the ExoMars
Aurora Programme for Solar System explora- rover to the surface of Mars, where it will search
tion decided to pursue the Mars exobiology for signs of life, past and present. The rover will
rover concept. In 2003, ESA issued a call for have the capability to drill to depths of 2 m to
instruments to be accommodated on the 2009 collect and analyze samples that have been
ExoMars rover. The ExoMars mission was shielded from the harsh conditions of the surface,
approved in 2005. A last-minute request to add where radiation and oxidants can destroy organic
a lander station resulted in a more complicated materials. The lander will be equipped with addi-
mission concept, requiring a larger launcher, tional instruments.
which could not be realized within the available ESA and Roscosmos have agreed to a well-
budget. A number of studies were necessary to balanced sharing of responsibilities for the differ-
redefine the mission, and thus the launch date ent mission elements. ESA will provide the TGO
was postponed, first to 2011 and then to 2013. and EDM in 2016 and the carrier and rover in
At the same time, NASA was experiencing cost 2018. Roscosmos will be in charge of the 2018
overrun problems with their ▶ Mars Science descent module and surface platform and will
Laboratory (MSL) mission that affected their furnish launchers for both missions. NASA con-
possibilities to independently realize a mission tributions to ExoMars include the TGO Electra
in 2016. In the course of 2009, ESA and NASA communications package and an important part
decided that they could accomplish more by of MOMA, the organic molecule characterization
joining forces. A scenario was defined for an instrument on the rover.
ESA-NASA joint program that had as ultimate
goal the realization of an international ▶ Mars
Sample Return (MSR) mission in the mid to late Overview
2020s. Within this program, ESA and NASA
defined the first two missions, in 2016 and Establishing whether life ever existed, or is still
2018. In late 2011, budget constraints in the active on Mars today, is one of the outstanding
US forced NASA to scale down its participa- scientific questions of our time. The ExoMars pro-
tion. ESA, NASA, and Roscosmos started a gram seeks to timely address this important scien-
dialogue aiming to realize the program as tific goal and to demonstrate key flight and in situ
a tripartite collaboration. However, in early enabling technologies underpinning European and
2012 NASA informed ESA and Roscosmos Russian ambitions for future exploration missions.
that they would not be able to contribute major The ExoMars scientific objectives are:
elements. Following a program reassessment
phase, in 2013 ESA and Roscosmos signed a • To search for signs of past and present life on
cooperation agreement to work in partnership Mars
792 ExoMars
• To investigate the water/geochemical envi- within the first billion years following planetary
ronment as a function of depth in the shallow formation. Conditions then were similar to those
subsurface when microbes gained a foothold on the young
• To study Martian atmospheric trace gases and Earth. Both planets had the necessary environ-
their sources mental conditions and ingredients for life,
namely, liquid water, carbon, and other essential
In support of these objectives, the following elements, as well as a source of energy. Life
technologies will be achieved: could have arisen in suitable locations, such as
in the vicinity of hydrothermal activity, where all
• Entry, Descent, and Landing (EDL) of a pay- requirements and ingredients could have existed,
load on the surface of Mars even if the standing bodies of water were ice
• Surface mobility with a rover covered. Not even intensive bombardment and
• Access to the subsurface to acquire samples possible volcanic resurfacing could have eradi-
• Sample acquisition, preparation, distribution, cated simple cells completely from the entire
and analysis planet’s surface. This marks Mars as a primary
target for the search for signs of life in our Solar
The ExoMars program includes missions to System.
the red planet for the 2016 and 2018 launch
opportunities (note that the orbits of Earth and The Martian Environment and the Need for
Mars are such that favorable opportunities occur Subsurface Exploration
about every 2 years to 26 months). For organisms to have emerged and evolved,
The 2016 TGO will accommodate scientific liquid water must have been present on Mars.
instruments for the detection of atmospheric trace Without it, most cellular metabolic processes
gases, the study of their temporal and spatial would not be possible. In the absence of water,
evolution, and the localization of their source life either ceases or slips into a quiescent mode.
regions. The EDM will contain engineering sen- Hence, the search for extinct or extant life auto-
sors to evaluate the lander’s descent performance matically translates into a search for liquid water-
and a small environmental station. rich environments, past or present.
The 2018 rover will carry a comprehensive The strategy to find traces of past biological
suite of instruments named after Louis Pasteur activity rests on the assumption that any surviv-
dedicated to exobiology and geochemistry ing signatures of interest will be preserved in the
research. The ExoMars Rover will travel several geological record, in the form of buried/encased
kilometers searching for signs of past and present remains, organic materials, and fossil communi-
life, collecting and analyzing samples from ties. Because current Martian surface conditions
within rocky outcrops and from the subsurface, are hostile to most known organisms, when
down to a depth of 2 m. looking for signs of extant life, the search meth-
In the case of a possible biosignature discovery, odology should focus on investigations in
confirmation of the results will require more thor- protected niches: in the subsurface and within
ough analyses than can be performed by remote surface outcrops. Therefore, the same sampling
robotic means. The ExoMars missions will contrib- device and instrumentation can adequately serve
ute immensely to determining what types of sam- both types of studies.
ples we should return to Earth for further studies. The rover’s surface mobility and the 2 m ver-
tical reach of the drill are both crucial for the
scientific success of the mission.
Basic Methodology The ExoMars rover will search for two types
of life-related signatures: morphological and
If life ever arose on the red planet, it probably did chemical. This will be complemented by an
when Mars was warmer and wetter, sometime accurate determination of the geological
ExoMars 793
context. Morphological information related to been processed by UV radiation, ionizing radia-

biological processes may be preserved on tion, and potential oxidants in the atmosphere
the surface of rocks. Possible examples include and on the surface of Mars. Any organic
the bio-mediated deposition of sediments, fos- biosignatures would be highly degraded in
silized bacterial mats, stromatolitic mounds, these samples.
etc. Essential for these studies are mobility For all the above reasons, the ExoMars drill
and an imaging system capable of covering will be able to penetrate and obtain samples from
from the meter scale down to submillimeter res- well-consolidated (hard) formations, at various
olution (to discern micro-textural information in depths, from the surface down to 2 m.
rocks). The successful NASA Mars Exploration
E
Effective chemical identification of bio- Rovers (MER) have demonstrated the past exis-
signatures requires access to well-preserved tence of wet environments on Mars using a
organic molecules. Because the Martian atmo- geologically oriented instrument package.
sphere is more tenuous than Earth’s, three impor- Their results have persuaded the scientific com-
tant physical agents reach the surface of Mars munity that mobility is a must-have require-
with adverse effects for the long-term preserva- ment for future missions. Recent discoveries
tion of organic compounds: (1) The ultraviolet from ESA’s ▶ Mars Express spacecraft have
(UV) radiation dose is higher than on our planet revealed multiple deposits containing salt and
and will quickly damage potential exposed clay minerals that can only form in the presence
organisms or biomolecules. (2) UV-induced pho- of liquid water. This reinforces the hypothesis
tochemistry is responsible for the production of that ancient Mars may have been wetter, and
reactive oxidant species that, when activated, can possibly warmer, than it is today. NASA’s Mars
also destroy organics; the diffusion of oxidants Science Laboratory (MSL) is presently studying
into the subsurface is not well characterized and surface geology and organics with the goal of
constitutes an important measurement that the identifying habitable environments in Gale cra-
mission must perform. (3) Finally, ionizing radi- ter. The 2018 ExoMars Rover constitutes the
ation penetrates into the uppermost meters of the next logical step in Mars exploration. It will
planet’s subsurface. This causes a slow degrada- have instruments to investigate whether life
tion process that, over many millions of years, ever arose on the red planet. It will also be the
can alter organic molecules beyond the detection first mission combining mobility and access to
sensitivity of analytical instruments. Please subsurface locations where organic molecules
note that the ionizing radiation effects are may be well-preserved, thus allowing for the
depth dependent: the material closer to the sur- first time to investigate Mars’ third dimension:
face is exposed to a higher dose than that buried depth. This alone is a guarantee that ExoMars
deeper. will break new scientific ground.
A major goal of ExoMars is to study ancient
(older than 3.5 billion years) sedimentary rock The Study of Martian Atmospheric Trace
formations and evaporitic deposits. However, Gases
the chemical record of early Martian life, if it Recent observations from the Planetary Fourier
ever existed, is likely to have escaped radiation Spectrometer (PFS) on ESA’s Mars Express and
and damage only if it is trapped in the subsurface from very high spectral resolution spectrometers
for long periods. Studies show that a subsurface using Earth-based telescopes have detected var-
penetration in the range of 2 m is necessary to iable amounts of methane in the atmosphere of
recover well-preserved organics from the very Mars. Based on photochemical models and on
early history of Mars. the current understanding of the composition of
Additionally, it is essential to avoid loose the Martian atmosphere, methane has a chemical
dust deposits distributed by eolian transport. lifetime of 300–600 years, which is very short
While driven by the wind, this material has on geological timescales. Thus, its presence
794 Exomoon
indicates a subsurface source that has recently ▶ Mars Express

(geologically speaking) released methane into ▶ Mars Sample Return Mission
the atmosphere. There are both geochemical ▶ Mars Science Laboratory
and biochemical processes that could produce ▶ Mars Stratigraphy
methane in the subsurface – its presence is not ▶ MER, Spirit and Opportunity (Mars)
sufficient to establish the nature of the source.
Current photochemical models cannot explain
the reported rapid space and time variations in
atmospheric methane concentration. Whether
Exomoon
geochemical or biochemical in origin, a confir-
mation of the methane observations would indi-
Nader Haghighipour
cate a dynamically active subsurface on Mars
today.
A scientifically exciting and credible experi-
ment can be conducted within the proposed mis-
sion concept of a 2016 science/telecom orbiter
delivering an EDL Demonstrator on direct entry Definition
to Mars (see next section). The scientific promise
of this mission is that it will reveal just how active An exomoon is a natural satellite (as opposed to
the Mars subsurface is, with the hope that it will an artificial satellite) of an extrasolar planet. In
explain the nature of that activity – geochemical analogy to the solar system where many of the
or biological. planets have moons around them, it is expected
that extrasolar planets will host moons as well.
Descent and Environment Science No exomoon has yet been discovered (as of
The 2016 mission scenario foresees the release of 2014), as the detection of exomoons is a very
the EDM from its hyperbolic arrival trajectory, complicated task with all current detection
approximately 3 days before touchdown. techniques.
Because this mission element constitutes a tech-
nological development rather than a science plat-
form, the nature of the sensors that it can
accommodate is relatively simple. The EDM
will rely on batteries to power a small environ- Exon
mental station after landing for a nominal surface
mission duration of 4 sols (Martian days). The Juli Peretó
science possibilities of such a mission are very Institut Cavanilles de Biodiversitat i Biologia
limited, but nevertheless useful. It will be possi- Evolutiva, Universitat de València, València,
ble to retrieve the atmospheric profile along the Spain
descent trajectory and obtain fundamental con-
straints for updating and validating the Mars stan-
dard atmospheric model.
Definition
In interrupted (split) ▶ genes, exon refers to each

See Also of the sequences in the primary transcript present
in the mature ▶ RNA sequence after removal of
▶ Habitability on Mars the ▶ introns (intervening sequences separating
▶ Mars the exons) by the splicing process.
Exoplanetary Atmospheric Retrieval 795
See Also transfer model coupled to an instrumental model

(taking into account spectral resolution, etc.).
▶ Gene (iii) Inverse model: This determines the best-
▶ Intron fitting state vector for the data by means of a
▶ RNA predetermined criterion, e.g., the chi-squared
metric, to determine the goodness of mock obser-
vations generated by F.
For astrobiology, two main objectives of
retrieval could be defined, the detection of habit-
Exoplanetary Atmospheric Retrieval
able conditions (related mainly to surface pres-
E
sure and surface temperature) and the detection
Philip von Paris
of bioindicators or biosignatures (e.g., CH4, O3,
etc.). Note, however, that retrieval alone does not
Universite de Bordeaux, Floirac, France
refer to the interpretation (biotic/abiotic) of
observed (or inferred) parameter values.
Key challenges for retrieval in an
exoplanetary context are (a) the small number
Definition
of data points to be expected, (b) the constraints
imposed by spectral resolution for currently
“Inversion method” or “retrieval” means deter-
planned instruments, and (c) the (most likely)
mining the (physical, chemical, etc.) characteris-
low signal-to-noise ratio of the observations.
tics of a system (state vector x) based on
Given these challenges, the state vector will
measurements (data vector d) obtained, for exam-
only contain very few variables and the model
ple, by telescope observations. The retrieved
M will be a simplified one. Even then, it appears
solution x0 for a given dataset d0 depends in gen-
difficult to overcome parameter degeneracies and
eral on the assumed model M of the particular
provide unique parameter solutions by means of
system and the data uncertainties. For exoplanets,
retrieval. For example, near-IR bands of H2O and
retrieval is possible only via spectrophotometric
CO2 overlap around 2 and 2.7 mm, or weak bands of
observations of the planetary atmosphere and
CO2 coincide with the strong O3 band around
surface. The spectrum is influenced by, for exam-
9.6 mm. Emission spectroscopy, which measures
ple, atmospheric composition and thermal struc-
the apparent temperature of a planet at a given
ture, clouds, and the scattering properties of the
wavelength, suffers from a degeneracy between
surface. This entry only deals with the retrieval of
the pressure level probed and the concentration of
atmospheric properties of rocky, potentially hab-
atmospheric constituents. For example, a measure-
itable exoplanets.
ment of a low apparent temperature could be due
to, for example, a low surface temperature (if the
atmosphere is transparent at the particular wave-
Overview length) or a high concentration of an atmospheric
species (and hence, the level probed is higher in the
Retrieval can be split into three parts. (i) System atmosphere where temperatures are lower).
model M: M calculates the state of the system Possible solutions for these challenges involve
based on the values of the state vector x. For this combining different types of observations that are
entry, M represents an atmospheric model that sensitive to different parameters (e.g., transmis-
calculates, for example, temperature and concen- sion and emission spectra). Furthermore, a priori
tration profiles. (ii) Forward model F: F simulates information should be applied, such as planetary
the dataset based on the results of M. In the case and orbital characteristics or detailed knowledge
of spectroscopic observations, F is a radiative of the host star.
796 Exoplanets, Discovery
sky) based on measurements of the changes in

Exoplanets, Discovery radial velocities using Doppler spectroscopy at
optical wavelengths that opened the exoplanet
David W. Latham1 and Nader Haghighipour2 floodgates. The conventional thinking was domi-
1
Harvard-Smithsonian Center for Astrophysics, nated by the architecture of the solar system, with
Cambridge, MA, USA the giant planets in wide circular orbits out past the
2
Institute for Astronomy, University of Hawaii- ▶ snow line, where conditions were cold enough
Manoa, Honolulu, Hawaii, HI, USA for them to form. The first radial-velocity orbit for
an unseen companion that could be called a giant
planet (if the orbit happened to be oriented nearly
Synonyms edge on) was published in 1989, but it defied
conventional wisdom on three counts. The orbital
Extrasolar planets period of only 84 days placed it far inside the snow
line, where conditions would have been too hot for
a gas giant to form; the orbit was eccentric, unlike
Definition any of the giant planets in the solar system, and the
mass was at least ten times that of Jupiter, which
Exoplanets, also known as extrasolar planets, are was uncomfortably large (and close to the mass of
planets beyond the solar system, orbiting around a small ▶ brown dwarf). Thus, most astronomers
stars other than the Sun. The discovery of the first dismissed the unseen companion of HD114762
exoplanet happened in 1992 when three planets (Latham et al. 1989) as a star or brown dwarf in
were discovered around the pulsar PSR 1257+12 an orbit seen nearly face on (thus minimizing the
(Wolszczan and Frail 1992). The first planet observed Doppler effect, which only measures the
around a Sunlike star (hence the name extrasolar velocity along the line of sight). It was assumed
planet) was discovered in 1995 by Mayor and that the observed orbital amplitude of 0.5 km/s was
Queloz. small because of projection effects and not because
the mass of the companion was small enough to be
a planet.
Overview In 1992 the detection of a system of unseen
planets orbiting the pulsar PSR B1257+12 was
The quest for planets beyond the solar system has announced (Wolszczan and Frail 1992). Unlike
achieved spectacular progress during the last the case of HD114762, these planets were detected
20 years. By the year 2015 astronomers may based on periodic variations in the arrival times of
well have discovered thousands of planets, the radio pulses from their rotating neutron host
many of which are similar to Earth, orbiting star. Follow-up observations, using the detection
Sunlike stars, some of which could possibly sup- of perturbations in the pulse arrival times consis-
port life as we know it. tent with gravitational interactions involving
After the discovery of the planetary system planetary-mass objects, soon demonstrated that
around pulsar PSR 1257+12 in 1992, the initial the masses were indeed small and similar to the
discoveries of extrasolar planets involved giant mass of Earth. Astronomers were puzzled how
planets with enough mass to induce measureable planets could survive the ▶ supernova event that
reflex motions in the stars they orbited, thus imply- created the neutron star or alternatively how they
ing the presence of unseen planets. Unlike what might have formed from the leftover debris, but it
many had anticipated that planets would be seemed clear that if planets could find a way to live
detected by astrometry through identifying around a pulsar, maybe they would prove to be
stellar motion in the plane of the sky and across common around normal stars similar to the Sun.
the line of sight, it was the stellar motions along the That is how the search for planets around Sun
line of sight (i.e., perpendicular to the plane of the like stars stood. Because our solar system was used
Exoplanets, Discovery 797
as a template, the search for giant planets (which the plane of the sky can in principle resolve this
could induce larger reflex motion on their host ambiguity because of the extra information pro-
stars) was focused on distances beyond the snow vided by the motion in two dimensions across the
line. It was in 1995 when an orbit for 51 Pegasi was line of sight. Indeed, in a few cases, astrometry has
announced that implied an unseen companion with provided orbital inclinations for radial-velocity
a mass similar to Jupiter and a remarkably short planets. However, the real breakthrough for deriv-
period of only 4.2 days (Mayor and Queloz 1995). ing actual masses for planets (and therefore
Soon after this announcement, many argued that confirming their planetary nature) came with the
because of the uncertainty in the inclination of the discovery of systems with orbits oriented close
planet’s orbit and therefore the value of its actual enough to be edge on as seen from the Earth so
E
mass, the unseen object may in fact be a more that the planets transited across the face of their
massive body such as a brown dwarf. The initial host stars. The first of such transiting planets was
resistance was, however, overwhelmed by the HD209458 (Charbonneau et al. 2000; Henry
announcement of several other low-Doppler- et al. 2000). Detection of planets using transit
amplitude orbits implying giant planets, often in photometry opened a new chapter in the
eccentric orbits close to their parent stars, some of exoplanetary science. Not only does this technique
them quite massive. Too many were being found allow the actual mass of the transiting planet to be
for them all to lie in orbits facing the Earth and thus determined (the orbital inclination with respect to
to possibly be brown dwarfs. A population of the plane of the sky is now known and is 90 ), it
extrasolar planets was being revealed, although also enables one to determine the radius of the
for any particular candidate the mass could not planet. The amount of stellar light blocked by the
be pinpointed until the inclination of the orbit to shadow of the planet allows the area, and therefore
the line of sight could be established. planet’s radius, to be measured. Knowing the mass
Ambitious Doppler surveys of hundreds and and radius of a planet, its bulk properties such as
even thousands of solar-type stars rapidly the density and surface gravity can be calculated.
expanded the known population of radial-velocity It turns out that transiting planets also provide
planet candidates. As the surveys were sustained a rich variety of opportunities to learn additional
over longer durations, planets with longer periods planetary astrophysics. This includes observa-
were discovered. As the instrumental techniques tions of planetary atmospheres during both the
improved and the velocity precisions improved, transits and the secondary eclipses (sometimes
the discoveries were extended to smaller masses called occultations), when the planet disappears
as well. As richer data sets were accumulated, behind its star, yielding the temperature and
systems with more than one planet were discov- chemical composition of the planet’s atmo-
ered. Finally, in the search for habitable planets, sphere. In some cases, the thermal emission
the surveys were focused on cool, low-mass stars. from the planet has been tracked as it moves
By now, a new class of bodies known as super- around in its orbit, thus revealing the changes in
Earths has been identified, some of which with temperature as more of the dayside comes into
masses only a few times that of Earth, orbiting in view and providing insights into the weather on
the habitable zones of their host stars. These the planet. The degree of the alignment of the
accomplishments are described in more detail in planet’s orbit with the equator of its host star can
the entry on ▶ Radial-Velocity Planets. also be determined in transiting system. Using the
To remove the ambiguity in the actual mass of ▶ Rossiter-McLaughlin effect, one can deter-
an unseen companion revealed by a radial-velocity mine the tilt of the planet orbit which can provide
orbit (traditionally termed as a ▶ spectroscopic important clues about the history of the formation
orbit by astronomers), the inclination of the orbit and evolution of the planet. Similar to planetary
to the line of sight must be established so that the systems detected by radial velocity, additional
observed orbital velocity can be corrected for pro- planets may exist in transiting systems (which
jection effects. Astrometry of the orbital motion on may or may not transit). These planets may reveal
798 Exoplanets, Modeling Giant Planets’ Atmospheres
their presence by perturbing the times at which Hubble Space Telescope. Earth-sized planets
transits of other planets are observed. Known as will be much more difficult to image and will
transit-timing variation (TTV) method, these var- require specialized space missions or ground-
iations have been used to detect several planets based “extremely large telescopes.”
by the Kepler space telescope. Perhaps surpris-
ingly, the detailed shape of a transit can pin down
the density of the host star, and if the shape is See Also
determined with sufficient precision, details such
as the oblateness of the planet due to rotation can ▶ 51 Pegasi B
also be determined. These topics are treated in ▶ Astrometric Planets
more detail in the entry ▶ Transiting Planets. ▶ Beta Pictoris b
An entirely different approach to the detection ▶ Coronagraphy
of unseen planets orbiting distant stars takes ▶ HD 209458b
advantage of an effect called gravitational ▶ Microlensing Planets
microlensing. When the line of sight to a distant ▶ Nulling Interferometry
star (the source) is intersected very precisely by an ▶ Pulsar Planets
intervening star (the lens), the light of the distant ▶ Radial-Velocity Planets
star is magnified by the lensing effect of the gravity ▶ Snow Line
of the intervening star. As the relative motion of ▶ Spectroscopic Orbit
the two stars changes the exact alignment, the ▶ Transiting Planets
changing magnification produces a light curve
that first brightens and then dims in a way that
can reveal information about the lensing star
(e.g., its mass). If a planet is orbiting the lens in Charbonneau D et al (2000) Detection of planetary transits
such a way that it passes close to the line of sight, it across a Sun-like star. Astrophys J Lett 529:45–48
can affect the brightening and introduce a feature Henry GW et al (2000) A transiting “51Peg-like” planet.
in the light curve. Although the precise alignment Astrophys J Lett 529:41–44
Latham DW et al (1989) The unseen companion of
needed to produce a microlensing event is very HD114762: a probable brown dwarf. Nature 339:38–40
rare, and the alignment to reveal an orbiting planet Mayor M, Queloz D (1995) A Jupiter-mass companion to
is even more rare, surveys to monitor hundreds of a solar-type star. Nature 378:355–359
millions of stars have detected and characterized Wolszczan A, Frail DA (1992) A planetary system around
the millisecond pulsar PSR1257 + 12.
several microlensing planets. These results and Nature 335:145–147
some details of the technique are described in the
entry ▶ Microlensing Planets.
An attractive way to study the characteristics
of an extrasolar planet would be to isolate its light Exoplanets, Modeling Giant Planets’
from that of the host star to produce a direct Atmospheres
image for further studies such as astrometry to
reveal the planet’s orbit or spectroscopy to Mark S. Marley
explore the structure and chemical composition NASA Ames Research Center, Moffett Field,
of its atmosphere. Such experiments present CA, USA
daunting technical challenges because of the
extreme brightness ratio and the tiny angular
separation between the planet and its host star, Keywords
even for the nearest systems. Direct images of a
few giant planet candidates in wide orbits around Atmosphere; Atmospheric structure; Planetary
hot stars have now been obtained both with spe- atmosphere; Reflection spectrum; Thermal
cial techniques from the ground and with the emission
Exoplanets, Modeling Giant Planets’ Atmospheres 799
Synonyms transparent and radiant energy escapes to space

and convection ceases. The pressure level in the
Giant planets, model atmospheres of atmosphere at which this radiative-convective
boundary lies depends upon the composition of
the atmosphere, the opacity of the major atmo-
Definition spheric constituents, gravity, and the tempera-
ture. For example, in the Earth’s atmosphere,
Models of the atmospheres of extrasolar giant the surface temperature is about 290 K. Only at
planets facilitate data interpretation and suggest a temperature of about 220 K near 100 mb (at the
new observations. tropopause) is the optical depth at the peak of the
E
Planck function low enough that the air can radi-
ate efficiently to space (Fig. 1). The tropopause
Overview lies at a few hundred millibars for the giant
planets as well (also Fig. 1).
Atmosphere models of extrasolar giant planets Meanwhile, incident flux from the primary
are crafted with the goal of understanding the star is either absorbed by the atmosphere or
vertical structure, composition, thermal profile, scattered back to space. Any atmosphere model
and dynamics of a given planet. Comparisons of must account for and conserve both the internal
model predictions of emitted and reflected flux to and this incident flux. To find a self-consistent
observed spectra provide insight into whether solution, a typical model atmosphere calculation
fundamental properties of the planet are begins with a draft temperature-pressure profile,
understood – or not. While there are several pos- which is then iterated upon until the energy flux
sible approaches to constructing an atmosphere through the atmosphere – accounting for depos-
model, the most traditional is forward modeling. ited incident flux – is conserved. During the iter-
Starting with a given set of assumptions, all of the ative process, the temperature-pressure profile
relevant equations governing the various physical must be adjusted such that the slope of the
properties affecting the atmosphere are model profile, dT/dP, does not exceed that
represented in a computer code, which predicts allowed by an adiabatic lapse rate. Profiles
the variation in atmospheric temperature and steeper than adiabatic initiate convection, which
composition with pressure in the atmosphere. In efficiently limits the maximum slope of the ther-
this entry, we consider the most important pro- mal profile. A final, self-consistent, profile is
cesses that must be accounted for in such models, known as a radiative-convective equilibrium pro-
as well as the ultimate goals of the modeling file. Samples of such model profiles for extrasolar
process. giant planets, including the computed convection
zones, are shown in Fig. 2. Further details on the
constraints that a model profile must satisfy are
Basic Methodology presented in the entry ▶ Atmosphere, Structure.
Since the radiative energy fluxes are the fun-
The most fundamental process to capture in a damental components of a model atmosphere
planetary atmosphere model is the transport of calculation, a necessary ingredient in any recipe
energy. In general, at depth in an atmosphere, for atmosphere model preparation is the knowl-
energy is transported by convection. For a giant edge of the atmospheric ▶ opacity. The opacity at
planet, this convection transports energy outward optical wavelengths must be known in order to
from the deep interior, as the planet cools slowly compute the amount of absorbed incident flux
over time. As heat is transported outward, the and the opacity at thermal wavelengths must be
atmosphere, which at depth is opaque, slowly known in order to calculate the emitted fluxes. In
becomes more transparent with increasing practice, this means the atmospheric opacity must
height. Eventually, the atmosphere becomes be known from the UV to the far infrared. There
0.0001
0.001
0.01
P (bar)
0.1
1 Jupiter
Saturn Earth
Uranus
10
50 100 150 200 250 300
T(K)
Exoplanets, Modeling Giant Planets’ Atmospheres, stratosphere – owing to the absorption of a fraction of
Fig. 1 Atmospheric temperature-pressure profiles for the incident ultraviolet light by photochemical products
Uranus, Saturn, Jupiter, and Earth (Neptune is similar to (ozone in the case of Earth, various hydrocarbon products
Uranus). In all four planets, the temperature increases with in the giants’ atmospheres). The upper boundary of the
depth below a few hundred millibars. In each planet, this troposphere is called the tropopause. Giant planet data are
atmospheric region – the troposphere – transports heat by from the Voyager Radio Science occultation experiments
convection from the deep interior, in the case of the giants, and the Earth profile is from the 1976 Standard Atmo-
or the surface in the case of Earth. In each atmosphere, the sphere (All data are available online http://atmos.nmsu.
temperature also rises at low pressure – the edu/planetary\_datasets/indextemppres.html)
are two required components: a calculation of at lower temperatures, the C would be found in
atmospheric composition and a calculation of CH4 and there would be somewhat more H2O
the opacity of each individual component. The present. At very cold temperatures, the gaseous
total atmospheric opacity is then the sum of the water would condense, leaving relatively little
contribution of each individual absorber, O in the gas phase. A rigorous chemical equilib-
weighted by its relative abundance. rium calculation is done for each atomic species
A determination of composition begins with a over the entire temperature and pressure range
calculation of which species would be present in under consideration. These calculations must
chemical equilibrium, given an assumed set of account for the rainout of species that condense.
initial atomic abundances. Typically, some mul- This means that once a species condenses, it
tiple of solar composition is assumed. The frac- does not stay in the atmosphere to react at
tion of elements heavier than H and He can be lower temperature but rather falls out by precip-
varied to determine the effects of sub- or super- itation in a gravity field. State-of-the-art chemi-
solar abundance. For any given assumed com- cal equilibrium calculations account for the
position, a unique abundance of molecules and interaction of thousands of different species
atoms can be calculated at any temperature and through a complex chemical network of
pressure, assuming chemical equilibrium. For reactions.
example, for a certain abundance of O and C, Once a gas composition has been computed
one would find that at high temperatures, these for a given temperature and pressure, the opacity
atoms are primarily found as CO and H2O, while of that gas must be computed as well. This
Exoplanets, Modeling
Giant Planets’ 0.001 0.02
Atmospheres, 0.05
Fig. 2 Pressure-
temperature profiles for 0.01 0.1
solar system-aged Jupiter- 0.2
like planets from 0.02 to
10 AU (left to right) from a 0.1 0.3
solar-type star. Thick lines 0.5
are convective regions P (bar)
while thin lines are 1.0 1.0
radiative regions. The 2.0
profiles at 5 and 10 AU E
10 3.0
show deviations that arise
from numerical noise in the 5.0
chemical equilibrium table
100 10
near condensation points
(Figure adapted from
Fortney et al. (2007))
1000
100 1000
T(K)
requires knowledge of the absorption spectrum of physics, can sometimes obscure underlying
atoms and molecules, often at high temperature. trends, so employing a variety of approaches is
In some cases, particularly for atoms and some often wise. An example of a simple cloud model
refractory molecules common in cool stars, opac- would be to assume a fixed particle size with
ities are well known. In other cases, particularly condensate distributed over some vertical thick-
for the molecules methane and ammonia, opaci- ness of atmosphere, typically one scale height.
ties are not well understood at high temperatures The total mass of cloud particles could be set
because of the difficulty in conducting appropri- equal to the total mass of condensible gas above
ate laboratory experiments. Furthermore, even the condensation level. Parameter space in such
for well-understood molecules, the effects of models can be easily explored by varying the
line broadening by the ubiquitous H2 molecules cloud thickness and particle size to understand
are not usually well known. the range of possible atmospheric responses.
In addition to the molecular opacity, the opac- A more complex model might attempt to com-
ity arising from condensate clouds must also be pute particle sizes and vertical condensate dis-
accounted for (▶ Clouds). Clouds can efficiently tribution given the computed or assumed vigor
scatter incident flux back to space, making a of atmospheric mixing. Highly complex
planet brighter in reflected light, and they can approaches could even attempt to model the
also trap infrared flux below. The former effect entire condensation process. While the latter
can cool the atmosphere while the latter results in approaches might be appropriate for studying a
warming. Any complete atmosphere model must known object, the simpler techniques can
thus attempt to model cloud formation. This is a quickly identify the range of planet brightnesses
difficult challenge, since cloud formation is a that might be expected, for example.
leading source of uncertainty even in highly Once all of these ingredients (composition,
sophisticated studies of Earth’s atmosphere. But opacity, cloud model) are in hand, an atmo-
the sizes and vertical distribution of cloud parti- spheric thermal profile for a given set of assumed
cles play leading roles in any model. conditions (interior heat flow, planet mass and
Approaches to cloud modeling can range gravity, distance from primary star, composition)
from the simple to the complex. Complex can be computed (Fig. 2). The overall thermal
models, while attempting to include more profile of an irradiated atmosphere depends both
upon the depth at which incident energy is mechanisms, heat up until the thermal emission
absorbed and internal sources of energy. Figure 2 from the layer equals the absorbed incident flux.
demonstrates that for planets more distant than a An inverted temperature structure with a warm,
few AU from their primary star, most incident radiative stratosphere overlying a cooler tropo-
energy is absorbed fairly deep in the atmosphere, pause is seen for several planetary atmospheres in
below the depth at which the atmosphere Fig. 1. In fact, almost all solar system planets with
becomes optically thick in the thermal infrared. an atmosphere exhibit a stratosphere. In Earth’s
This is because most gases are more transparent atmosphere, ozone absorbs ultraviolet (UV) light,
in the optical than in the infrared. As a result, the which warms the stratosphere to 270 K, about
absorbed incident energy simply adds to the inter- 50 K warmer than the temperature at the top of
nal energy being transported outward by convec- the troposphere. Solar system giant planet atmo-
tion and the temperature profile resembles that of spheres are heated by UV absorption by a com-
Jupiter shown in Fig. 1. bination of methane and hydrocarbon
For those giant planets found closer to their photochemical products, including C2H2 and
primary stars, the radiative-convective boundary C2H6, and photochemically produced hazes.
is deeper but absorption of incident flux still The atmosphere of Jupiter (Fig. 1) provides a
occurs at a similar altitude (to the extent that specific example. Without an energy source, the
composition is unchanged). Thus, the large inci- planet’s middle atmosphere would be close to
dent flux upon a hot Jupiter is absorbed above the 104 K (the skin temperature for Jupiter with
radiative-convective boundary (Fig. 2). As a Teff = 124 K), as seen above the tropopause in
consequence, an isothermal layer appears Fig. 1. In the region where most of the incident
between the top of the deep convective zone UV flux is absorbed (near 10 mbar), there is little
and the region of the atmosphere in which inci- overlap between a 100 K Planck function and the
dent flux is absorbed. In this case, the deep important thermal opacity sources, so little flux
internal heat flow is distinct from the thermal- can be emitted. Since the absorbed energy cannot
ized incident radiation and the global tempera- be radiated away by a 100 K atmosphere, the
ture distribution is no longer relatively atmosphere warms and the Planck function
homogeneous and equator to pole temperature moves to shorter wavelengths. Eventually, the
gradients can be large. blue side of the Planck function overlaps the
In the lower pressure region lying above the strong n4 methane fundamental vibrational band
radiative-convective boundary (and generally and the n9 ethane bands at 7.7 and 12.2 mm,
above thick clouds), the atmosphere is in radia- allowing the atmosphere to radiatively cool,
tive equilibrium and – lacking any source of balancing the absorbed incident flux. As in other
turbulence – generally stable, hence the name solar system giant planet atmospheres, these
stratosphere. In this region, an idealized gray strong mid-infrared bands of ethane and methane
gas atmosphere would become isothermal at the act as a thermostat, regulating the stratospheric
skin temperature T0 = (1/2)1/4 Teff, where Teff is temperatures.
the effective temperature (the temperature of a Models of exoplanet atmospheres must like-
blackbody radiating the same total energy as the wise account for gases which may be important
actual planet). However, a real, non-gray, atmo- absorbers or emitters in the upper atmosphere.
sphere can be opaque to incident radiation over Important species may be those expected in
some spectral range at low pressures and simul- chemical equilibrium, for example, TiO in the
taneously be relatively transparent at infrared hottest atmospheres, or photochemical products.
wavelengths. In this case, more incident energy To account for the latter requires modeling com-
may be absorbed than can be emitted by an iso- plex reaction chains that can be triggered by the
thermal atmosphere with temperature T0. As a absorption of incident UV flux and adds yet
result, the atmospheric layer with strong absorp- another challenge to constructing complete atmo-
tion must, in the absence of other energy transport spheric models.
Exoplanets, Modeling 0.8

Giant Planets’
Atmospheres, Rayleigh scattering
0.7
Fig. 3 Observed
geometric albedo of Uranus Uranus Lambert sphere
and Jupiter compared to 0.6
one possible model of HD
Geometric albedo
209458b and the upper 0.5 Jupiter
limit from the
Microvariability and
0.4
Oscillations of STars space
telescope (MOST). The
geometric albedo of a deep 0.3 E
Rayleigh scattering HD 209458b
atmosphere and a 0.2
Lambertian sphere are
shown as well
0.1 Most
0.0
0.4 0.5 0.6 0.7 0.8 0.9 1.0
Wavelength (mm)
Key Research Findings the red). At shorter wavelengths, photochemi-

cally produced hazes on Jupiter absorb blue
Atmosphere models resulting from the tech- light, while Uranus’ relatively clear atmosphere
niques discussed in the previous section have a efficiently scatters incident flux. HD 209458b,
number of applications. Their predictions can which lacks any bright clouds and hosts multiple
assist efforts to detect and characterize planets strong absorbing molecules, is almost black
and, when applied to specific known planets, beyond 0.5 mm. Figure 4 illustrates model spectra
can identify important processes actually acting for giant planets at a range of separations from a
on exoplanet atmospheres. A complete review of Sun-like star. Once water clouds form beyond
both types of applications is well beyond the about 2 AU, the planet brightens notably at red
scope of this entry. Instead, here we present a and longer wavelengths as can be seen from com-
few indicative research findings. For a detailed parison of the dotted and solid lines.
discussion of specific model predictions or planet In the future, space-based coronagraphs will
interpretations, the relevant literature should be directly image planets in reflected light at optical
consulted. wavelengths. For such observations, an under-
standing of exoplanet spectra and color will be
Planets in Reflected Light needed. An example illustrates the need: given an
The influence of the various physical processes orbital separation from the primary star, a single
discussed above can be found in reflected and photometric detection, combined with an
transmitted light and thermal emission observa- assumed phase function (relative brightness as a
tions of extrasolar planets. For example, Fig. 3 function of the angle star-planet-observer) and
compares observed ▶ albedo spectra of Jupiter bounds placed on the geometric albedo, would
and Uranus with a computed spectrum of the allow a crude estimate of the planet’s size.
hot Jupiter HD 209458 b. Jupiter’s relative Assuming an upper limit geometric albedo less
brightness at longer wavelengths arises from its than 0.75 (the pure Rayleigh scattering limit) and
bright, thick ammonia clouds (which scatter inci- a lower limit of 0.06 (typical of low-albedo aster-
dent light efficiently) and ten times lower abun- oids), for example, would result in an uncertainty
dance of methane (which absorbs efficiently in in the radius inferred for a directly imaged planet
Exoplanets, Modeling 0.8

Giant Planets’
Atmospheres, 3 AU
Fig. 4 Model geometric
albedo spectra for a Jupiter- 2 AU
0.6
mass planet at various
distances from a solar-type
Geometric albedo
1.0 AU
star. Dotted lines are
models for the 3 and 2 AU
planets but without cloud 0.4 0.7 AU
opacity (Modified from
Cahoy et al. (2010)) 0.5 AU
0.2 AU
0.2
0.0
0.4 0.5 0.6 0.7 0.8 0.9 1.0
Wavelength (mm)
of a factor of 3.5. A bright planet with a radius (because of methane absorption in the red)
slightly larger than Earth’s could not be distin- than cloudy, cooler planets. Reflectivity in the
guished from a dark planet with Neptune’s radius blue is further influenced by stratospheric
on the basis of brightness alone. If the planet were hazes produced by photochemical processes.
also detected by other means, for example, radial Thus, discerning planet characteristics from
velocity or astrometric methods, then the known color alone will be challenging. Spectra,
mass would discriminate between these two even low-resolution spectra, as shown in Figs. 3
extremes. Without such detection, however, the and 4, will be much more informative, if
nature of the planet would have to be discerned obtainable.
by spectroscopic or photometric methods. Even
low-resolution spectroscopy likely will be Understanding Thermal Emission
beyond the reach of modest aperture space- The spectrum of any planet is composed of two
based coronagraphic telescopes. This means that components: scattered radiation incident from the
planets will have to be characterized, at least planet’s star and thermal emitted flux from the
initially, by their broadband colors. planet. The thermal flux represents both energy
Indeed based on our experience in the solar arising from processes interior to the planet and
system, broadband colors of giant planets at first reradiated absorbed incident radiation. For solar
seem to be promising markers for discerning system planets, these two components of the
planet type (Fig. 5 for the giants). Uranus and spectrum are usually well separated in wave-
Neptune are blue, while Jupiter and Saturn are length, but for the hottest exoplanets, there can
red. However, as also shown in Fig. 5, model be substantial overlap between thermal radiation
calculations of planet color reveal that color, as and scattered incident light. For a planet, such as
does a set of spectra, depends sensitively on the a transiting planet, with a known radius, the ther-
presence or absence of clouds and atmospheric mal emission spectrum is often equated for con-
composition as well as viewing geometry. The venience, wavelength by wavelength, to the
right-hand panel of the figure shows model thermal emission from a ▶ blackbody. For an
planet colors for a variety of star-planet separa- isothermal solid sphere with emissivity unity,
tions, planet masses, and compositions. Cloud- the observed spectrum would equal that of a
less planets, regardless of mass, are much bluer blackbody with a fixed temperature. However,
a 0.6 b 0.6
0.4 0.4
0.2 0.2
0 0
V–R
V–R
−0.2 −0.2
−0.4 −0.4 E
Jupiter
−0.6 Neptune −0.6
Saturn
−0.8 Uranus −0.8
Titan
−1 −1
−1.5 −1 −0.5 0 0.5 −1.5 −1 −0.5 0 0.5
R−1 R−1
Exoplanets, Modeling Giant Planets’ Atmospheres, and cyan denote Jupiters with one and three times solar
Fig. 5 Color-color plot (see ▶ Color Index; V, R, and abundance of heavy elements and Neptunes with 30 times
I refer to wavelength intervals defined by standard astro- solar abundance (respectively). Fading dot intensity
nomical filters in the visual, ref, and near-infrared portions denotes color change as viewing angle varies from 0 to
of the spectrum) in (V-R) vs. (R-I) showing observed 180 in 10 increments (Modified from Cahoy
location of solar system giant planets and Titan (a, left, et al. (2010))
stars) as well as model planets (b, right, dots). Red, blue,
for a real planet the flux will differ from that of a for a monotonically increasing temperature pro-
blackbody with the same radius, and at each file with depth, means higher brightness temper-
wavelength a “brightness” temperature TB(l) ature. High-opacity spectral regions correspond
may be defined (equal to the temperature of a to lower pressures and lower temperatures. How-
blackbody radiating the observed intensity at a ever, if there is an inverted temperature profile,
specific wavelength l). Although thermal emis- then a situation could emerge where
sion data for transiting planets are often reported TB(l1) > TB(l2) and TB(l2) < TB(l3) where
in terms of TB(l), such data must be regarded T(l1) > T(l2) > T(l3). This is a commonplace
with some care. Except in special cases (e.g., an occurrence in the atmospheres of solar system
isothermal atmosphere), brightness temperature giants and the chromospheres of stars.
is not a measure of physical temperature or effec- Furthermore, because of limb (edge) effects,
tive temperature. Rather it gives a weighted mea- this range in pressures from which flux emerges
sure of atmospheric temperatures over a range of also varies over the disk. Thus, even for a gray
pressures from which flux emerges from the atmosphere, with constant optical depth as a
planet. function of wavelength, the brightness tempera-
Crudely, the brightness temperature can ture is in general not equal to the effective tem-
indeed be equated to the temperature in the atmo- perature at all wavelengths. For these reasons,
spheric region which most contributes to the while brightness temperatures are useful short-
emergent flux. However, since atmospheric opac- hands to convey information about planetary
ity can vary dramatically with wavelength, the spectra, they must be regarded with some caution.
brightness temperature can vary substantially For example, 8 mm Spitzer observation of the hot
with wavelength. In regions of low opacity, flux Neptune GJ 436b yields a brightness temperature
emerges from deeper in the atmosphere which, of 712 36 K which is modestly above the
predicted effective temperature. Since we do not quest to understand the processes responsible for
expect, in general, for TB = Teff, the information the observed heating.
content of this single data point is limited. With
atmosphere models and additional data points, Disequilibrium Chemistry
the value of each brightness temperature mea- As noted above, the first approach to constructing
surement increases. an exoplanet atmosphere model is to assume
chemical equilibrium. However, an important
Hot Stratospheres limitation to conventional one-dimensional
One of the more intriguing observational findings models of mean atmospheric structure is the
of the ▶ hot Jupiters has been the ubiquity of hot neglect of vertical mixing. Vertical transport
stratospheres. Observations primarily by the plays an important role when the dynamical time-
Spitzer Space Telescope have ascertained that scale is short compared to a particular chemical
as many as half of the transiting hot Jupiters’ equilibrium timescale, as is the case for CO in the
sport hot stratospheres, emitting at temperatures atmosphere of Jupiter. While methane is the most
well in excess of the skin temperature. As in the abundant C-bearing molecule in Jupiter’s visible
solar system Jovian atmospheres discussed in the atmosphere, in the deep atmosphere, where tem-
previous section, in exoplanet atmospheres, a peratures are higher, the abundance of CO is
balance must be struck between the absorption substantially larger. Since the C-O bond is very
of incident radiation and thermal emission. For strong, the conversion time from CO to CH4 in a
hot Jupiters, which are so warm that even the parcel of rising gas is correspondingly long. As a
most refractory Ti- and V-bearing compounds consequence, vertical mixing through the atmo-
do not condense, TiO and VO gases may be sphere transports CO from the deep atmosphere
exceptionally important absorbers. These gases, to the visible atmosphere where it can be
while not abundant, have extraordinarily large detected.
absorption cross sections across the entire optical Evidence for CO mixing ratios enhanced over
spectrum. When present, these molecules can that expected in chemical equilibrium is widely
absorb much of the remarkably high incident noted in the brown dwarf literature and tentative
flux at altitudes above 1 mbar, where the atmo- evidence is apparent in the available exoplanet
sphere is optically thin in the thermal infrared. data as well. The study of disequilibrium pro-
The atmosphere, therefore, becomes very hot, as cesses is a classic success of the traditional for-
hot as 2,000 K or more, which is hot enough for ward modeling approach to understanding
emission by the near-infrared and optical bands planetary atmospheres. Models that assume pure
of water, CO, and even TiO to balance this influx chemical equilibrium fail to match available data
of energy. (spectra and photometry of exoplanets), which
Discerning which extrasolar planets possess leads to a search for additional physical mecha-
hot stratospheres has become a major endeavor nisms to consider. When an additional important
for exoplanet science, particularly when utilizing physical process is accounted for in the models,
the IRAC instrument aboard the Spitzer Space the predictions compare more favorably with data
Telescope. It seems clear that the TiO/VO mech- and knowledge has been gained.
anism is plausible for some of the hottest extra-
solar giant planets, but likely is not the full Dynamics
explanation for cooler exoplanets with hot strato- Atmosphere models can also be used to study
spheres. It is likely that an as yet unknown atmospheric dynamics on a global scale. The
absorber, perhaps one produced by disequilib- same models that compute mean, global condi-
rium or photochemical processes, is responsible tions can also be adapted to compute profiles at
for stratospheric heating in some fraction of the specific points on a planet. Coupled with three-
hot Jupiters. The techniques of forward modeling dimensional fluid flow computations, the nature
discussed here will continue to be required in the of the atmospheric circulation of giant planets,
accounting for energy transport and rotation, can system. This is because molecular species that
be computed. The atmospheric redistribution of condense below the Jovian clouds (e.g., H2O,
energy by winds is unquestionably of paramount H2S, NH3) and thus are protected from photodis-
importance for the hot Jupiters, and the efficiency sociation will be gaseous in such hot atmo-
of redistribution controls the global temperature spheres. Some of these species, such as H2S, are
map and consequently the phase variation of easily photodissociated and will likely produce
thermal emission, which has been successfully new or unexpected species. Sulfur and nitrogen
measured for multiple planets. Since a planet’s compounds, in particular, may be important
thermal emission can arise from different depths players in hot Jupiter photochemistry and perhaps
in the atmosphere at different wavelengths, any haze production. While the carbon photochemis-
E
variation in redistribution efficiency with altitude try has been studied, recent work on photochem-
will manifest itself as differing thermal emission istry in water-bearing H2He atmospheres
maps as a function of wavelength. Ultimately, suggests that compounds including CO2, HCN,
coupled models of radiative transfer and dynam- and C2H6 will be present well in excess of the
ics, similar to terrestrial global circulation abundance predicted by equilibrium chemistry.
models, will be required to understand all of the As a consequence, CH4 may be underabundant,
contributing factors. as has indeed been found for the atmosphere of
one exoplanet. Photochemical products may also
play a role in the formation of hot stratospheres.
Future Directions This area is certainly rich for further study.
Second, ever more sophisticated global circu-
As more and better data become available for the lation modeling of atmospheric dynamics will
transiting extrasolar giant planets, particularly become increasingly important. Current models
from the James Webb Space Telescope, the are hampered by the need to compute radiation
opportunities for more new and interesting balance and dynamics through a three-
science – and the requirements for ever more dimensional model atmosphere over tens of thou-
sophisticated models – will grow. Ultimately sands of time steps. Such calculations are a time-
direct imaging of cool, mature planets in reflected consuming process even for current generation of
light will join the nascent field of direct imaging computers. Continuing advances in computa-
of young, luminous giant planets at the forefront tional speed and improvements in the computa-
of exoplanet science. In the nearer term, a number tion and handling of molecular opacities will
of new directions can be expected from theoreti- permit more sophisticated and, hopefully, more
cal models of giant exoplanet atmospheres. realistic model cases to be considered.
First, atmospheric molecules can be dissoci- Finally, of course, a great variety of
ated by the absorption of ultraviolet light, a pro- exoplanets will likely be found. The variations
cess that happens high in the atmosphere before in planet atmospheric composition, gravity, tem-
most incident UV light is scattered back to space. perature, dynamics, photochemistry, and as yet
Photochemical products can then participate in unrecognized processes will certainly be great
complex reaction chains, producing various enough to attract the full attention of the next
molecular products. A familiar example is atmo- generation of atmospheric modelers.
spheric ozone in Earth’s stratosphere, which ulti-
mately results from the photodissociation of
molecular oxygen. Photochemical products can See Also
themselves become important players in the
atmospheric radiative transfer of giant planets. ▶ Adiabatic Processes
Photochemistry has long been expected to be ▶ Albedo
important for hot Jupiter atmospheres and will ▶ Atmosphere, Model 1D
likely be far more complex than in the solar ▶ Atmosphere, Structure
808 Exopolymers
▶ Atmosphere, Temperature Inversion Rowe JF et al (2008) The very low albedo of an extrasolar
▶ Biomarkers, Spectral planet: MOST space-based photometry of HD 209458.
Astrophys J 689:1345
▶ Clouds Spiegel DS, Silverio K, Burrows A (2009) Can TiO
▶ Color Index explain thermal inversions in the upper atmospheres
▶ GCM of irradiated giant planets? Astrophys J 699:1487
▶ Grey Gas Model Stevenson KB et al (2010) Possible thermochemical dis-
equilibrium in the atmosphere of the exoplanet GJ
▶ Habitability of the Solar System 436b. Nature 464:1161
▶ Habitable Planet, Characterization
▶ Hot Jupiters
▶ Mini-Neptunes
▶ Non-Grey Gas Model: Real Gas Atmospheres
▶ Rayleigh Scattering Exopolymers
Lucas J. Stal
References and Further Reading Department of Marine Microbiology, Royal
Netherlands Institute of Sea Research (NIOZ),
Burrows A, Hubbard WB, Lunine JI, Liebert J (2001) The Yerseke, The Netherlands
theory of brown dwarfs and extrasolar giant planets.
Rev Mod Phys 73:719
Cahoy K, Marley M, Fortney J (2010) Exoplanet albedo
spectra and colors as a function of planet phase, sepa- Keywords
ration, and metallicity. Astrophys J 724:189
Chamberlain J, Hunten D (1987) Theory of planetary
atmospheres. Academic, Orlando Biofilm; Cyanobacteria; Diatoms; Microalgae;
Deming D, Seager S, Richardson LJ, Harrington J (2005) Polymers
Infrared radiation from an extrasolar planet. Nature
434:740
Fortney JJ, Marley MS, Barnes JW (2007) Planetary radii
across five orders of magnitude in mass and stellar Synonyms
insolation: application to transits. Astrophys
J 659:1661–1672 EPS; Extracellular polymeric substances; Extra-
Fortney JJ, Lodders K, Marley MS, Freedman RS cellular polymers; Extracellular polysaccharides
(2008) A unified theory for the atmospheres of the
hot and very hot Jupiters: two classes of irradiated
atmospheres. Astrophys J 678:1419
Freedman RS, Marley MS, Lodders K (2008) Line and Definition
mean opacities for ultracool dwarfs and extrasolar
planets. Astrophys J Suppl 174:504
Hubeny I, Burrows A, Sudarsky D (2003) A possible Exopolymers are polymers that are deposited
bifurcation in atmospheres of strongly irradiated stars outside the (microbial) cell. These polymers are
and planets. Astrophys J 594:1011 predominantly composed of carbohydrates, but
Knutson HA, Charbonneau D, Allen LE, Burrows A, many contain various other components such as
Megeath ST (2008) The 3.6–8.0 0 mm broadband
emission spectrum of HD 209458b: evidence for an proteins, DNA, and glycolipids. Some
atmospheric temperature inversion. Astrophys exopolymers consist of neutral sugars such as
J 673:526 glucose, while others are acidic in nature and
Lodders K, Fegley B (2002) Atmospheric chemistry in contain a variety of charged groups such as the
giant planets, brown dwarfs, and low-mass dwarf stars.
I. Carbon, nitrogen, and oxygen. Icarus 155:393 uronic acids, carboxy groups, sulfated sugars, or
Marley MS, Gelino C, Stephens D, Lunine JI, Freedman pyruvate groups. The molecular structure and
R (1999) Reflected spectra and albedos of extrasolar composition of exopolymers is therefore highly
giant planets. I. Clear and cloudy atmospheres. diverse and complex. Exopolymers form the
Astrophys J 513:879
Marley MS, Fortney J, Seager S, Barman T (2007) In: matrix of biofilms in which the microorganisms
Reipurth B, Jewitt D, Keil K (eds) Protostars and are embedded but also serve a plethora of other
planets V. University of Arizona Press, Tucson, 733 p functions.
Exopolymers 809
Overview mineral particles, resulting in sediments with a

high erosion threshold. Hence, exopolymers
Exopolymers are by definition polymers that are have found application in sediment stabilization.
deposited outside the cell wall. These include Exopolymers may bind calcium and magnesium
sheaths or investments in which the cell, but ions, thereby controlling calcification. It may act
often also aggregates or chains or threads of cells, as an anti-calcification agent, but it may also
are wrapped and represent a more or less structural allow local calcium carbonate precipitation and
part of the organism. Such sheaths may be highly hence the tertiary structure of the polymer may
structured and may have a typical morphology, determine the morphology of the calcium
while other infestations are more diffuse. Other carbonate. This is the case with the calcium
E
exopolymers do not have a close association with carbonate platelets known as coccoliths, which
the cells that produce them. They are exuded into are produced by certain marine microalgae.
the environment and “dissolve” as colloidal Exopolymers protect the microorganisms from
exopolymers in the water or form in the benthic being grazed and immobilize (heavy) metals
environment the biofilm matrix in which the and other toxic compounds and antibiotics. The
microorganisms are embedded. latter is an important problem for curing infec-
Exopolymers that are exuded into the environ- tions by pathogenic bacteria, while the former
ment are usually the result of unbalanced growth, has found application in bioremediation of pol-
for instance, when photosynthetic organisms fix luted soils. Many exopolymers may form a gel
CO2 but are limited by nutrients. In that case, the through the absorption of large amounts of water.
fixed carbon cannot be accommodated by the syn- This gel protects the organism from desiccation.
thesis of structural cell material and will be depos- Exopolymers can also scavenge important nutri-
ited as polysaccharide. Part of it will be stored ents from the environment, thereby providing the
intracellularly as a reserve compound but the biofilm organisms with an important advantage
bulk will find its way as colloidal exopolymers. compared to free-living pelagic microorganisms.
The same may be the case with chemotrophic Finally, exopolymers can act as flocculants.
microorganisms that live at the expense of Some benthic cyanobacteria have been shown
energy-rich but nutrient-poor resources. to produce it in order to bind mineral particles,
Exopolymers can also exude as a result of the thereby clearing in the water column above them.
gliding motility by some bacteria, notably by
▶ cyanobacteria. The exopolymers are exuded
through specific pores in the cell wall and interact
See Also
with the substrate on which the organism glides,
leaving behind a trail of exopolymer. The mecha-
▶ Archean Traces of Life
nism of gliding motility is still poorly understood.
▶ Biomarkers, Morphological
Exopolymers are not just waste products.
▶ Cyanobacteria
They may serve a variety of important functions
for the organism that produces them as well as for
the ecosystem as a whole. It has been shown that References and Further Reading
benthic diatoms metabolize the exuded
exopolymers as a carbon and energy source in Braissant O, Decho AW, Przekop KM, Gallagher KL,
Glunk C, Dupraz C, Visscher PT (2009) Characteristics
the dark. Exopolymers form the matrix of and turnover of exopolymeric substances in a
biofilms. They may be important for attachment hypersaline microbial mat. FEMS Microbiol Ecol
of microorganisms to surfaces. This may be 67:293–307
important for the success of the organism but Branda SS, Vik Å, Friedman L, Kolter R (2005) Biofilms:
the matrix revisited. Trends Microbiol 13:20–26
represents also a problem known as fouling.
Decho AW (1990) Microbial exopolymer secretions in
Exopolymers form coherent networks and ocean environments: their role(s) in food webs and
matrixes that may interact covalently with marine processes. Oceanogr Mar Biol 28:73–153
810 Exothermic
Hoagland KD, Rosowski JR, Gretz MR, Roemer SC

(1993) Diatom extracellular polymeric substances – Exozodiacal Light
function, fine structure, chemistry, and physiology.
J Phycol 29:537–566
Hoiczyk E, Hansel A (2000) Cyanobacterial cell walls: Daniel Rouan
news from an unusual prokaryotic envelope. LESIA, Observatoire Paris-Site de Meudon,
J Bacteriol 182:1191–1199 Meudon, France
Pereira S, Zille A, Micheletti E, Moradas-Ferreira P, De
Philippis R, Tamagnini P (2009) Complexity of
cyanobacterial exopolysaccharides: composition,
structures, inducing factors and putative genes Keywords
involved in their biosynthesis and assembly. FEMS
Microbiol Rev 33:917–941
Stal LJ (2003) Microphytobenthos, their extracellular Dust particles; Exoplanet; Interplanetary dust;
polymeric substances, and the morphogenesis of inter- Scattering
tidal sediments. Geomicrobiol J 20:463–478
Sutherland IW (2001) Biofilm exopolysaccharides: a
strong and sticky framework. Microbiology 147:3–9
Synonyms
Exozodi
Exothermic
Jacques Reisse Definition

Université Libre de Bruxelles, Brussels, Belgium
The exozodiacal light is the analog in extrasolar
planetary systems of the ▶ zodiacal light seen in
Definition the solar system. It corresponds to the light emit-
ted by the central star, scattered – or reemitted in
The term exothermic refers to a reaction or a the infrared – by interplanetary small dust parti-
physical transformation which releases energy if cles concentrated in the mean orbital planetary
it takes place at constant volume or which releases plane, that is, the local ▶ ecliptic. It is often
▶ enthalpy if it takes place at constant pressure. abbreviated as exozodi. This may be a strong
A familiar exothermic reaction is the ▶ combus- emission that can prevent direct detection of
tion of coal. The enthalpy change (DH) during an planets belonging to the system.
exothermic reaction or transformation taking place
at constant pressure is negative by definition. The
enthalpy release takes the form of a heat release. Overview
In the solar system, the zodiacal light appears as a

flame-shaped glow just above the horizon, visible
See Also to the naked eye in good atmospheric conditions
and when the moon has not risen. It is seen only
▶ Combustion toward the west after sunset or in the east before
▶ Endothermic sunrise and is aligned along the zodiacal belt,
▶ Enthalpy hence its name. It is due to sunlight reflected by
dust particles of 1–100 mm in size, concentrated in
the plane of the ▶ ecliptic, mainly in its central
region (approximately up to the asteroid belt).
Exozodi Because the dust particles are heated by the sun-
light to a temperature comparable to the Earth’s
▶ Exozodiacal Light (300 K), they also emit in the infrared. It is
Experimental Evolution 811
Exozodiacal Light,
Fig. 1 HST coronagraphic
image of the star Fomalhaut
at 0.6 mm, showing together
the dust belt responsible for
the exozodiacal light
emission and the planet
Fomalhaut b (white square)
just within the inner
boundary of the dust belt, at
two different dates
generally admitted that extrasolar planetary sys- been identified as exhibiting an exozodiacal light,
tems must harbor a similar content of dust and thus at least in the infrared domain: Beta Pictoris and
exhibit an analog of the zodiacal light in the visible 51 Ophiuchus are two examples.
and infrared. The origin of this dust can be prim-
itive material in the ▶ protoplanetary disk, but
more often in mature systems, it corresponds to See Also
debris left after planet formation when the
remaining ▶ planetesimals suffered collisions ▶ Debris Disk
and evaporation. The question of the intensity of ▶ Exoplanets, Discovery
the exozodiacal light is extremely important when ▶ Zodiacal Light
dealing with the direct detection and characteriza-
tion of planets in the ▶ habitable zone, because References and Further Reading
exozodiacal light intensity may be so large that it
would dominate by several orders of magnitude Cockell CS, Herbst T, Léger A, Absil O, Beichman C,
the signal from a planet. In the solar system, for Benz W, Brack A, Chazelas B et al (2009) Darwin an
instance, the zodiacal light is 300 times stronger experimental astronomy mission to search for extraso-
lar planet. Exp Astron 23:435
than the emission of the Earth at a wavelength of Kalas P, Graham JR, Chiang R et al (2008) Optical images
10 mm. The name exozodi was proposed as an of an Exosolar planet 25 light years from Earth.
abbreviation when its importance was realized Science 322:1345
and discussed in its details. It is generally assumed Reach WT (1997) The structured zodiacal light: IRAS,
COBE, and ISO observations, diffuse infrared radiation
that the detection of an earthlike planet should be and the IRTS. ASP conference series no. 124, pp 33–40
possible, provided that the exozodi does not exceed Smith BA, Terrile RJ (1984) A circumstellar disk around
ten times the solar system value. A consequence is beta pictoris. Science 226:1421–1424
that before launching an expensive space mission Stark CC, Kuchner MJ, Traub WA, Monnier JD,
Eugene S, Mark C, Chris K, Bertrand M et al (2009)
that aims at direct detection of exoplanets, precur- 51 Ophiuchus: a possible beta pictoris analog mea-
sor missions or possibly measurements from the sured with the Keck interferometer Nuller. Astrophys
ground should assess the relative frequency of large J 703(2):1188–1197
and medium exozodi intensities in the visible and
in the infrared. Figure 1 shows an image of the
exosystem Fomalhaut where the exozodi is
resolved as a ring and contributes to the back- Experimental Evolution
ground against which a planet has been directly
detected (Kalas et al. 2008). A few stars have ▶ Evolution, In Vitro
812 Explosive Nucleosynthesis
Synonyms
Explosive Nucleosynthesis
EXPOSE-E; EXPOSE-R; EXPOSE-R2
▶ Nucleosynthesis, Explosive
EXPOSE Definition
Hervé Cottin EXPOSE-E, EXPOSE-R, and EXPOSE-R2

Laboratoire Interuniversitaire des Systèmes are exposure facilities developed by ESA in
Atmosphériques, Université Paris Est-Créteil, the 2000s to investigate the effect of space envi-
Créteil, France ronment (especially energetic radiations) on
various chemical and biological samples.
EXPOSE-E was installed on the ▶ International
Keywords Space Station outside of the European Colum-
bus module on the European Technology Expo-
Exposure facility; International Space Station; sure Facility (EuTEF), from February 2008 to
Low Earth orbit; Photobiology; Photochemistry; August 2009, while EXPOSE-R was installed
Space radiations outside of the Russian module Zvezda
EXPOSE, Fig. 1 The EXPOSE facility (480 520 the experiments, one layer of samples is exposed to space,
327.5 mm) is made of three experiment trays into while another layer is fitted just below the first one to be
which four square sample carriers (77 77 26 mm) used as flight controls
are fitted (Credit Kayser-Threde – Germany). In most of
EXPOSE 813
EXPOSE, Table 1 Astrobiology experiments selected after the first call of ESA in 1996 and finally accommodated on
EXPOSE-R
Experiment Topic of research Principal investigator
AMINO Photochemical processing of organic compounds, including H. Cottin, LISA/UPEC – Créteil
amino acids, RNA fragments, and samples relevant to (France)
cometary and Titan chemistry
ORGANIC Study of the evolution of organic matter in space (PAHs) P. Ehrenfreund – Leiden
Observatory (NL)
ROSE-1/ Study of the impact of extraterrestrial UV radiation on C. Cockell – Open University
ENDO microbial primary producers (algae, cyanobacteria) (UK)
ROSE-2/ Study of the protective effects of osmophilic microorganisms R. Mancinelli – SETI Institute,
OSMO enclosed within gypsum-halite crusts NASA Ames (USA) E
ROSE-3/ Study of the protection of spores by meteorite material against G. Horneck – DLR (Germany)
SPORES with space conditions: UV, vacuum, and ionizing radiation/
R3D radiation dosimetry
ROSE-4/ Study of the photoproducts resulting from exposure of dry J. Cadet – CEA Grenoble
PHOTO DNA samples or bacterial spores to solar UV radiation (France)
ROSE-5/ Study of the mutational spectra of Bacillus subtilis spores N. Munakata – University of
SUBTIL induced by space vacuum and/or solar UV radiation Tokyo (Japan)
ROSE-8/PUR Study of the biologically effective dose of solar extraterrestrial G. Rontó – Research Lab. for
UV radiation by biological dosimetry Biophysics, Budapest (Hungary)
EXPOSE, Table 2 Astrobiology experiments selected after the second call of ESA in 2004 and finally accommodated
on EXPOSE-E
ADAPT Study of molecular adaptation strategies of microorganisms to P. Rettberg – DLR (Germany)
different space and planetary UV climate conditions
DOSIS/ Passive radiation dosimetry at the sample sites G. Reitz – DLR
DOBIES (Germany)/F. Vanhavere – SCK
CEN (Belgium)
LIFE Study of the resistance of lichens and lithic fungi at space S. Onofri – Università degli studi
conditions della Tuscia di Viterbo (Italy)
PROCESS Study of photochemical organic chemistry relevant to comets, H. Cottin, LISA/UPEC – Créteil
meteorites, Mars, and Titan (France)
PROTECT Study on the resistance of spacecraft isolates to outer space for G. Horneck – DLR (Germany)
planetary protection purposes
R3D-2 Active radiation dosimetry (VIS, UV-A, UV-B, UV-C; LET D-P. Häder – University of
spectra of cosmic radiation) Erlangen (Germany)
SEEDS Study of plant seed as a terrestrial model for a Panspermia D. Tepfer – CNRS, Versailles
vehicle and as a source of universal UV screens (France)
from March 2009 to March 2011. EXPOSE-R2 Overview

is a reload of EXPOSE-R with new experiments
which departed to the space station in July 2014, A typical EXPOSE facility is presented in Fig. 1.
and should return to Earth early 2016. Data from six temperature sensors connected to
814 Exposed Surface Bioburden
EXPOSE, Table 3 Astrobiology experiments selected by ESA to be accommodated on EXPOSE-R2

BIOMEX Study of the stability of pigments, cellular components, and J.P. de Vera – DLR
extremophiles under space and Mars-like conditions (Germany)
BOSS Study of the stability of bacteria exposed as biofilms and microbial mat P. Rettberg – DLR
communities under space and Mars-like conditions (Germany)
PSS Study of the stability of organic compounds with a prebiotic relevance H. Cottin, LISA/
under space and Mars-like conditions UPEC – Créteil (France)
the three trays, four UV-B sensors, and one radi-

ometer are recorded every 10 s. Exposed Surface Bioburden
Experiments selected by ESA to be
implemented on EXPOSE-E, EXPOSE-R, and Catharine A. Conley
EXPOSE-R2 are presented in Tables 1, 2 (from NASA Headquarters, Washington, DC, USA
Rabbow et al. 2009), and 3.
Definition
See Also The term “exposed surface ▶ bioburden” is used

to indicate the number of viable ▶ microorgan-
▶ BIOPAN isms that are carried on internal and external
▶ International Space Station spacecraft surfaces that are available for particu-
▶ Ionizing Radiation, Biological Effects late and gas exchange, and from which microor-
▶ Lyman Alpha ganisms could reach a planetary environment
▶ Photolysis following the nominal landing of a spacecraft.
▶ Solar UV Radiation, Biological Effects
▶ Space Environment See Also
▶ UV Radiation
▶ Bioburden
▶ Microorganism
Cottin H et al (2008) Heterogeneous solid/gas chemistry

of organic compounds related to comets, meteorites, EXPOSE-E
Titan and Mars: in laboratory and in lower Earth orbit
experiments. Adv Space Res 42:2019–2035
Rabbow E et al (2009) EXPOSE, an astrobiological ▶ EXPOSE
exposure facility on the International Space Station –
from proposal to flight. Orig Life Evol Biosph
39:581–598
Rabbow E et al (2012) EXPOSE-E, an ESA Astro- EXPOSE-R
biology mission 1.5 years in space. Astrobiology
12:374–386 ▶ EXPOSE
Exposure Facilities 815
likelihood of viable interplanetary transfer of

EXPOSE-R2 life and the limiting factors of ▶ Panspermia.
First attempts were already performed during
▶ EXPOSE the Gemini program (Hotchin et al. 1968); they
were continued in a more sophisticated manner
during the Apollo program and up to the present
studies on the ▶ International Space Station (ISS)
Exposure Facilities (Horneck et al. 2010).
Gerda Horneck
E
DLR German Aerospace Center, Institute of Overview
Germany To expose chemical or biological samples to
outer space or selected parameters of this extreme
environment, several exposure facilities were
Keywords developed for attachment to the outer shell of
spacecraft (Table 1).
Outer space parameters; Photochemical reac- The first sophisticated exposure device was
tions; Space experiments; Survival built in 1972 by ▶ NASA, the Microbial Ecology
Evaluation Device (▶ MEED) for the Apollo
16 mission (Taylor et al. 1974). MEED was
Synonyms mounted to the distal end of the TV boom of the
Command Module during the extravehicular
Exposure platforms; Exposure trays activity phase of the trans-Earth coast. It was
composed of 798 sample cuvettes with quartz
windows as optical filters with the optional pro-
Definition vision of ventilation holes for access to space
vacuum (Fig. 1). Using a solar positioning
Exposure facilities are technical devices attached device, MEED was oriented directly perpendicu-
to the outer shell of a spacecraft with the purpose lar to the sun. In 1983, Spacelab 1 (SL 1) carried
to expose ▶ organic molecules, ▶ microorgan- the exposure facility ES029 in its cargo bay
isms, and other small biological systems to (Horneck et al. 1984). ES029 consisted of an
outer space or to selected parameters of this exposure tray partitioned in four square quartz-
extreme environment. covered compartments (Fig. 2). Two of the com-
partments were vented to the outside, allowing
access to space vacuum. The other two compart-
History ments were hermetically sealed with a constant
pressure of 105 Pa. Each compartment accommo-
Soon after the advent of space flight, the accessi- dated 79 dry samples in the upper layer, allowing
bility of space has been used to study specific UV exposure, and the same number was kept in
questions of astrobiology, such as the role of the bottom layer as flight dark controls.
stellar ▶ UV radiation in the evolution of poten- UV-irradiated samples were placed beneath an
tial precursors of life, the role of the UV radiation optical filtering system composed of interference
climate in prebiotic and biological evolution on filters for narrow wavebands (220 nm, 240 nm,
Earth or any other celestial body, and the 260 nm, and 280 nm) and neutral density filters.
816 Exposure Facilities
Exposure Facilities, Table 1 Exposure facilities used on space missions to study the stability of organic molecules
and the survival of biological systems in outer space (Horneck et al. 2010)
Mission Mission Exposure Exposure
Year name characteristics facility duration Space parameter studied
1966 Gemini IX Earth orbit Collecting/ 16 h 47 min Space, solar UV
and XII (300 km alt) exposure (GIX)
device 6 h 24 min
(GXII)
1972 Apollo 16 Lunar mission MEED Vacuum: 1 h Space vacuum; solar UV:
20 min, UV: 254, 280 nm
10 min
1983 Spacelab 1 Earth orbit ES029 Vacuum: Space vacuum; solar UV: >170;
(240 km alt.) 9 days, UV: 220; 240; 260; 280
19 min–5 h
17.5 min
1984–1990 LDEF Earth orbit Exostack 2,107 days Space vacuum; solar UV
(500 km alt.)
1992–1993 EURECA Earth orbit, ERA 327 days Space vacuum; solar UV: >110;
sun pointing >170; >280; >295; 220; 230;
260; 290 nm
1993 Spacelab D2 Earth orbit RD- Vacuum: Space vacuum; solar UV: 190;
UVRAD 10 days; UV: 210; 220; 230; 260; 280; >190;
5–120 min >304; >313; >314; >315; >316;
>317 nm
1994 Foton 9 Earth orbit Biopan-1 14.8 days Space vacuum; solar UV
1997 Foton 11 Earth orbit Biopan-2 10 days Space vacuum. solar UV
1999 Foton 12 Earth orbit Biopan-3 12.7 days Space vacuum. solar UV
1999 MIR-Perseus Earth orbit Exobiologie 98 days Space vacuum, solar UV
1999 Terrier Black Ballistic flight SERTIS 395 s Space vacuum, solar EUV
Brant Rocket <304 km alt. (30.4 nm)
2004 Terrier Mark Ballistic flight 350 s High speed atmospheric entry
70-Improved <304 km alt.
Rocket
2005 Foton-M2 Earth orbit Biopan-5 14.6 days Space vacuum; solar UV: >170;
>280; >320; >400 nm
Stone-5 Meteorite entry in Earth’s
atmosphere
2007 Foton-M3 Earth orbit Biopan-6 10 days Space vacuum, solar UV: > 110;
>200; > 290; > 400 nm
Stone-6 Meteorite entry in Earth’s
atmosphere
2008–2009 ISS-EuTeF Earth orbit EXPOSE-E 1.5 years Space vacuum; solar
UV: > 110 nm; simulated Martian
atmosphere and UV
climate: > 200 nm
2009–2011 ISS Earth orbit EXPOSE-R 2 year Space vacuum, solar
UV: > 110; > 200 nm
A nontransparent shutter with optical windows samples were exposed to space vacuum for
was used to achieve precise irradiation intervals 10 days and for predefined periods (from 19 min
during the “hot phase” of the mission, when dur- to 5 h 17.5 min) to solar UV radiation. The tem-
ing several orbits the cargo bay of the shuttle was perature ranged from 17 C to 35 C, the highest
perpendicularly pointing towards the sun. The values occurring during the “hot phase” of the
Exposure Facilities 817
Exposure Facilities,
Fig. 1 Exposure facility
MEED, which was
mounted on the camera
beam of the lunar orbiter of
the Apollo 16 mission
(Credit: NASA)
Exposure Facilities, Fig. 2 Exposure tray of the ES029

experiment, which was mounted on a cold plate in the
cargo bay of SL 1
Exposure Facilities, Fig. 3 Exposure tray of the Exobi-
ology Radiation Assembly (ERA), which was mounted on
the EURECA platform (Credit: ESA, from Horneck
mission. A similar device was flown in 1993 with et al. 2010)
the experiment UVRAD during the German SL
D2 mission that provided a “hot phase” during
two orbits at the end of the mission. (1984–1990), was achieved during the LDEF
Long-term exposures of organic chemical mission within the German experiment Exostack
compounds and microorganisms to space started (Horneck et al. 1994). Frequent opportunities for
with the NASA ▶ Long Duration Exposure Facil- short-duration exposure experiments (up to
ity (LDEF) and were continued with the Euro- 15 days) of molecules and biological systems
pean Retrievable Carrier (EURECA) mission were provided by ESA’s Biopan facilities, cylin-
(Fig. 3) (Innocenti and Mesland 1995), and the drical pan-shaped containers with a deployable
French PERSEUS mission on the Russian MIR lid mounted on the outer surface of the descent
station (Rettberg et al. 2002). The longest expo- module of a Russian Foton satellite (Fig. 4)
sure of microorganisms to space, about 6 years (Demets et al. 2005). The Foton satellite was
818 Exposure Facilities
Fig. 4 Biopan facility with
lid opened, which was
mounted on the outside of
Foton satellites (Credit
ESA, from Horneck
et al. 2010)
Fig. 5 EXPOSE-E facility
(arrow), which was
mounted on the EuTef
platform of the ESA
Columbus facility of the
ISS (Credit ESA)
also used to study the mineral decomposition and trays and of ERA. The EXPOSE-E facility was
microbial survival during atmospheric reentry mounted during extravehicular activity to the
within the ▶ STONE experiments (Brandstätter European Columbus Module of the ISS as part
et al. 2008; Cockell et al. 2007; de la Torre of the European Technology Facility (EuTeF)
et al. 2010). platform in February 2008 and retrieved in Sep-
Advanced exposure facilities with up to four tember 2009. Experiments on prebiotic chemical
times the capacity of the ES029 experiment of SL evolution were located in one tray of EXPOSE-E;
1 were developed by the ▶ European Space the other two trays accommodated different
Agency (ESA) with the Exobiology Radiation microbial systems, either exposed to outer space
Assembly (ERA) for the ▶ EURECA mission conditions (space vacuum and solar UV spectrum
(Fig. 3) and the ▶ EXPOSE facilities attached to of l >110 nm) or to simulated Mars surface
the ISS (Rabbow et al. 2009). One EXPOSE unit climate (600 Pa pressure, 95 % CO2, and solar
consists of three trays (Fig. 5), each housing four UV of l >200 nm). The second EXPOSE facil-
compartments similar to those of the SL exposure ity, EXPOSE-R, was launched in November
Exposure Trays 819
2008 and will remain attached to the URM-D transfer of photosynthesis: an experimental demon-
platform, an external ISS facility at the Russian stration of a selective dispersal filter in planetary island
biogeography. Astrobiology 7:1–9
Zvezda module, for about 2 years. EXPOSE-E de la Torre R, Sancho LG, Horneck G, de los Rı́os A,
and EXPOSE-R house a total of 13 different Wierzchos J, Olsson-Francis K, Cockell CS,
experiments that are performed in international Rettberg P, Berger T, de Vera J-PP, Ott S, Martinez
cooperation (Baglioni et al. 2007). Before Frı́as J, Melendi PG, Lucas MM, Reina M, Pintado A,
Demets R (2010) Survival of lichens and bacteria
launching into space, all EXPOSE experiments exposed to outer space conditions – results of the
were tested in carefully designed ground simula- Lithopanspermia experiments. Icarus 208:735–748
tion experiments and in an experiment sequence Demets R, Schulte W, Baglioni P (2005) The past, present
test using the Planetary and Space Simulation and future of Biopan. Adv Space Res 36:311–316
Horneck G, B€ ucker H, Reitz G, Requardt H, Dose K, E
Facilities (PSI) at the ▶ German Aerospace Cen- Martens KD, Mennigmann HD, Weber P (1984)
ter DLR. The EXPOSE experiments will be con- Microorganisms in the space environment. Science
tinued with EXPOSE-R2. 225:226–228
Horneck G, B€ ucker H, Reitz G (1994) Long-term survival
of bacterial spores in space. Adv Space Res
14(10):41–45
Horneck G, Klaus DM, Mancinelli RL (2010) Space
See Also
microbiology. Microbiol Mol Biol Rev 74:121–156
Hotchin J, Lorenz P, Hemenway C (1968) The survival of
▶ BIOPAN terrestrial microorganisms in space at orbital altitudes
▶ Exobiologie Experiment during Gemini satellite experiments. Life Sci Space
Res 6:108–114
▶ EXPOSE
Innocenti L, Mesland DAM (eds) (1995) EURECA scien-
▶ Extreme Environment tific results. Adv Space Res 16(8):1–140
▶ Foton Capsule, Spacecraft Rabbow E, Horneck G, Rettberg P, Schott JU, Panitz C,
▶ Inactivation L’Afflitto A, von Heise-Rotenburg R, Willnecker R,
Baglioni P, Hatton J, Dettmann J, Demets R, Reitz
▶ International Space Station
G (2009) EXPOSE, an astrobiological exposure facil-
▶ Lichens ity on the International Space Station – from proposal
▶ Lithopanspermia to flight. Orig Life Evol Biosph 39:581–598
▶ Long Duration Exposure Facility Rettberg P, Eschweiler U, Strauch K, Reitz G, Horneck G,
Wänke H, Brack A, Barbier B (2002) Survival
▶ Mars
of microorganisms in space protected by meteorite
▶ MEED material: results of the experiment EXOBIOLOGIE
▶ Panspermia of the PERSEUS mission. Adv Space Res 30:
▶ Spore 1539–1545
Taylor GR, Spizizen J, Foster BG, Volz PA, B€ ucker H,
▶ STONE
Simmonds RC, Heimpel AM, Benton EV (1974)
A descriptive analysis of the Apollo 16 microbial
response to space environment experiment. Bioscience
References and Further Reading 24:505–511
Baglioni P, Sabbatini M, Horneck G (2007) Astrobiology

experiments in low Earth orbit: facilities, instrumenta-
tion, and results. In: Horneck G, Rettberg P (eds)
Complete course in astrobiology. Wiley-VCH, Ber-
lin/New York, pp 273–320 Exposure Platforms
Brandstätter F, Brack A, Baglioni P, Cockell CS,
Demets R, ME EH, Kurat G, Osinski GR, Pillinger ▶ Exposure Facilities
JM, Roten C-A, Sancisi-Frey S (2008) Mineraogical
alteration of artificial meteorites during atmospheric
entry. The STONE-5 experiment. Planet Space Sci
56:976–984
Cockell CS, Brack A, Wynn-Williams DD, Baglioni P,
Brandstätter F, Demets R, Edwards HME, Gronstal
Exposure Trays
AL, Kurat G, Lee P, Osinski GR, Pearce DA, Pillinger
JM, Roten C-A, Sancisi-Frey S (2007) Interplanetary ▶ Exposure Facilities
820 Extended Red Emission
Extended Red Emission Extinct Radionuclides
William M. Irvine Daniele L. Pinti

University of Massachusetts, Amherst, MA, USA GEOTOP Research Center for Geochemistry and
Synonyms
ERE Definition
Extinct radionuclides are atoms with unstable

nuclei that have undergone radioactive decay
Definition
into atoms of another element in a geologically
short time. These elements, which originated
The broad emission band observed in some inter-
from primordial processes of stellar
stellar environments (e.g., ▶ reflection nebulae,
nucleogenesis in supernovas, are now extinct in
▶ planetary nebulae, ▶ HII regions, interstellar
nature because of their short half-lives of 100 Ma
“▶ cirrus clouds”) extending between about
or less, relative to the age of the Earth (4.5 Ga).
600 and 800 nm is referred to as extended red
An example is 26Al, with a half-life of 0.7 Ma,
emission (ERE). It is thought to be the result of
which decays into 26Mg. These radionuclides
photoluminescence from ▶ interstellar dust,
allow dating of objects formed early in the history
although it is not clear whether the carrier is
of the Solar System. Indeed these objects contain
carbon-rich or silicate-rich material. The spatial
traces of the daughter elements produced by
distribution of ERE is, in many cases, similar to
decay of the extinct radionuclides incorporated
that of ▶ polycyclic aromatic hydrocarbons
prior to their extinction. Intense decay heat gen-
(PAHs).
erated by these radionuclides is thought to have
allowed the total melting of medium- to small-
size planetary bodies, driving early planetary
See Also
differentiation.
▶ Cirrus Cloud
▶ HII Region
▶ Planetary Nebula See Also
▶ Polycyclic Aromatic Hydrocarbon
▶ Reflection Nebula ▶ Differentiation, Planetary
▶ Earth, Age of
▶ Geochronology
References and Further Reading ▶ Magnetic Fields and Planetary Systems
Formation
Whittet DCB (2003) Dust in the galactic environment, ▶ Radiogenic Isotopes
2nd edn. Institute of Physics, Philadelphia
Extensive Plain Extinction Event
▶ Vastitas, Vastitates ▶ Mass Extinctions

Extraterrestrial Delivery of Organic Compounds 821
Extinction, Interstellar or Extinction-Level Event

Atmospheric
▶ Mass Extinctions
Daniel Rouan
Meudon, France
Extracellular Polymeric Substances
▶ Exopolymers
Definition E
The extinction is the decrease of the light inten-

sity of a celestial object due to scattering and/or Extracellular Polymers
absorption by an intervening medium. The
medium can be of telluric origin (molecules, ▶ Exopolymers
aerosol in the atmosphere) or astrophysical
(small dust particles in the ▶ interstellar
medium); in the later case one speaks of inter-
Extracellular Polysaccharides
stellar extinction. Extinction is measured in
▶ magnitudes and the wavelength at which it
▶ Exopolymers
applies should be indicated. The interstellar
extinction is larger in the blue than in the red
and becomes less and less important as the wave-
length increases in the infrared domain. Because Extrachromosomal Genetic Element
of this differential effect, extinction is responsi-
ble for the so-called reddening of distant stars. ▶ Plasmid
Reddening is characterized by the ▶ color
excess. The radio domain is essentially not
affected by extinction. Interstellar extinction is
generally noted AF, where F is the symbol of Extrasolar Planets
the wavelength considered (U, B, V, etc.),
defined by a passband filter. When no indication ▶ Exoplanets, Discovery
is given, it is assumed that the extinction is in the
visible band (V). Typical extinction by the
average ▶ interstellar medium in the plane of
our Galaxy is AV = 2 mag/kpc (1 kpc is about
Extraterrestrial Delivery of Organic
Compounds
3,000 light-years).
André Brack
Centre de Biophysique Moléculaire CNRS,
See Also Orléans Cedex 2, France
▶ Color Index
▶ Dust Grain Keywords
▶ Interstellar Medium Amino acids; Chirality; Comets; Impacts; Mete-
▶ Magnitude orites; Micrometeorites; Prebiotic compounds;
▶ Reddening, Interstellar Space experiments
822 Extraterrestrial Delivery of Organic Compounds
Definition detected in Comet Hyakutake in 1996, including

ammonia, methane, acetylene (ethyne), acetoni-
The primitive Earth experienced a large spectrum trile (methyl cyanide), and hydrogen isocyanide.
of impactors ranging from the huge Mars-sized In addition, the study of the Hale-Bopp comet in
impactor which created the Moon to cosmic dust 1997 led to the detection of methane, acetylene,
less than 1 mm in size. A great number of organic formic acid, acetonitrile, hydrogen isocyanide,
molecules, including amino acids, have been isocyanic acid, cyanoacetylene, formamide, and
found in carbonaceous chondrites. Micrometeor- thioformaldehyde.
ite collection and analysis from the Greenland The Stardust mission collected samples of
and Antarctic ice sheets suggest that the Earth Comet Wild 2 and returned them to Earth in
accreted large amounts of extraterrestrial com- January 2006 for laboratory analysis. Unexpect-
plex organic molecules. Intense bombardment edly, most of the comet’s rocky matter formed
probably caused some chemical reprocessing of inside the solar system at extremely high temper-
the Earth’s primitive atmosphere. Laboratory and ature. The grains contain a variety of organic
space experiments support plausibility of the functional groups (alcohol, ketone, aldehyde,
extraterrestrial delivery of organics to the primi- carboxylic acid, amide, nitrile). The protein-
tive Earth. building amino acid glycine has also been
discovered.
Comets orbit on unstable trajectories and
Overview sometimes collide with planets. The collision of
Comet Shoemaker-Levy 9 with Jupiter in July
Delivery of Extraterrestrial Organic Matter 1994 gave a recent example of such events.
Such collisions were probably more frequent
Comets 4 billion years ago, the comets orbiting around
Comets are the richest planetary objects in the Sun being more numerous. Comets may
organic compounds known so far. Ground-based therefore be an important source of organic mol-
observations have detected hydrogen cyanide and ecules delivered to the primitive Earth
formaldehyde in the coma of comets. In 1986, (Ehrenfreund and Charnley 2000; Despois and
on-board analyses performed by the two Russian Cottin 2005). However, it is unlikely that whole
missions Vega 1 et 2, as well as observations comets could have safely delivered organics to
obtained by the European mission Giotto and the Earth. They exploded either while crossing
the two Japanese missions Suisei and Sakigake, the atmosphere or when impacting the Earth’s
demonstrated that Halley’s comet contains subsurface.
stantial amounts of organic material. On average, The most detailed study of a comet ever
dust particles ejected from the Comet Halley attempted is about to be performed by the ESA
nucleus contain 14 % of organic carbon by Rosetta robotic spacecraft launched in March
mass. About 30 % of cometary grains are domi- 2004 which reached the comet 67P/
nated by light elements C, H, O, and N, and 35 % Churyumov–Gerasimenko in August 2014. The
are close in composition to the carbon-rich mete- spacecraft consists of two main elements: the
orites. The presence of organic molecules, such Rosetta space probe orbiter, which features
as purines, pyrimidines, and formaldehyde poly- twelve instruments, and the Philae robotic lander,
mers, has also been inferred from the fragments with an additional nine instruments (Glassmeir
analyzed by the Giotto PICCA and Vega PUMA et al. 2007).
mass spectrometers. However, there was no
direct identification of the complex organic mol- Meteorites
ecules probably present in the cosmic dust grains Carbonaceous chondrites delivered organic
and in the cometary nucleus. Many chemical materials to the early Earth. They contain from
species of interest for exobiology have been 1.5 % to 4 % of carbon, for the most part as
organic materials. One hundred kilograms of the NMR, applied to the organic fraction of Murchi-
Murchison meteorite, a CM2 type carbonaceous son extracted under mild conditions allowed to
chondrite that fell in Australia in 1969, have extend its indigenous chemical diversity to tens
been extensively analyzed (Pizzarello 2007; of thousands of different molecular compositions
Pizzarello and Shock 2010, and references and likely millions of diverse structures (Schmitt-
therein). Murchison organic materials are gen- Kopplin et al. 2010).
erally classified according to their solubility in Most of the amino acids detected in the carbo-
water and organic solvents. Insoluble and solu- naceous chondrites are chiral but present as race-
ble components represent, respectively, 70 % mate, i.e., the L- and D-enantiomers are present in
and 30 % of total carbon components. The insol- equal proportions. However, Cronin and
E
uble organic material is referred as kerogen-like, Pizzarello (1997) found L-enantiomer excesses
a poorly identified insoluble macromolecular (e.e.) in six a-methyl-a-amino acids from the
material of complex composition with average Murchison (e.e. 2.8–9.2 %) and Murray (e.e.
elemental formula C100H46N10O15S4.5. NMR, 1.0–6.0 %) carbonaceous chondrites. An enantio-
IR, and pyrolysis analyses suggest the presence meric excess up to 18 % has been measured for
of aromatic ring clusters bridged by aliphatic isovaline, 2-methyl-2-aminobutyric acid. These
chains, with peripheral branching and functional amino acids (isovaline, 2-amino-2,3-
groups. The insoluble organic material releases a dimethylpentanoic, a-methyl norvaline,
variety of aromatic and heteroatomic hydrocar- a-methyl valine, and a-methyl norleucine) are
bons as well as a suite of alkyl dicarboxylic acids either unknown or rare in the terrestrial biosphere
up to C18 chain length under conditions similar and cannot therefore be attributed to terrestrial
to those of hydrothermal vents (Yabuta contamination (Pizzarello 2007). In addition, the
et al. 2007). indigeneity of D- and L-isovaline enantiomers is
The soluble organic compounds of the Mur- supported by carbon and hydrogen isotopic data
chison meteorite represent a diverse and abun- (Pizzarello et al. 2003; Pizzarello and Huang
dant group of organics that vary from small 2005).
water-soluble compounds such as amino acids Several organic and inorganic phases of the
and polyols up to 30 carbon-long hydrocarbons. carbonaceous chondrite matrix were separated
This diversity has been analyzed in detail for the and subjected to the Soai reaction, i.e., the addi-
amino acid. The total number of meteoritic tion reaction of i-Pr2Zn to pyrimidine-5-
amino acids is about 100. All the possible carbaldehyde. Asymmetric autocatalysis with
a-amino acids up to seven carbon were identified amplification of chirality gave pyrimidyl alkanol
as well as large abundances of N-substituted, with enantiomeric excesses of detectable level.
cyclic, b-, g-, d-, and e-amino acids. Eight The asymmetry resides in powders after extrac-
protein-building amino acids (glycine, alanine, tion with water and solvents as well as in the
proline, leucine, isoleucine, valine, aspartic acid, insoluble organic material obtained after demin-
and glutamic acid) have been found. Nucleic acid eralization. Asymmetry is not found any longer in
bases, purines and pyrimidines, have also been the insoluble organic matter after hydrothermal
found in the Murchison meteorite (Stoks and treatment and in meteorite powders from which
Schwartz 1982). No ribose, the sugar moiety all organics had been removed (Kawasaki
which links together the nucleic acid building et al. 2006).
blocks, was detected in meteorites. Vesicle- The meteoritic enantiomeric excesses may
forming fatty acids have been extracted from help to explain the emergence of a homochiral
different carbonaceous meteorites (Deamer (one-handed) life. Each amino acid, with the
1985, 1998). exception of glycine, exists in two enantiomeric
A combination of high-resolution analytical forms, L and D, but proteins use only the L ones.
methods, composed of organic structural spec- Proteins adopt asymmetric rigid geometries,
troscopy FTICR/MS, UPLC-QTOF-MS, and a-helices and b-sheets, which play a key role in
the catalytic activity. Homochirality is now per year. This amazing dominance of micromete-
believed to be not just a consequence of life but orites already suggests their possible role in deliv-
also a prerequisite for life, because stereoregular ering complex organics to the early Earth
structures such as protein b-sheets, for example, 4.2–3.9 Ga ago when the micrometeorite flux
do not form with mixtures of monomers of both was probably enhanced by several orders of mag-
handednesses. The use of one-handed nitude. Antarctic micrometeorite flux measure-
biomonomers also sharpens the sequence infor- ments suggest that a huge mass (5 1024 g)
mation of the biopolymers. For a polymer made of micrometeorites was accreted by the Earth
of n units, the number of sequence combinations during the first 300 Ma of the post-lunar period.
will be divided by 2n when the system uses only At least, 20 wt% of the micrometeorites
homochiral monomers. Taking into account the survive unmelted by atmospheric entry. As their
fact that enzyme chains are generally made of kerogen fraction represents about 2.5 wt% of
hundreds of monomers, the tremendous gain in carbon, this amounts to a total mass of kerogen
simplicity offered by the use of monomers of 2.5 1022 g on the early Earth’s surface
restricted to one handedness is self-evident. equivalent to a 40-m-thick global layer
The excess of L-amino acids found in the Mur- (Maurette and Brack 2006). This delivery repre-
chison meteorite may result from the processing sents more carbon than that present in the bio-
of the organic mantles of interstellar grains by mass of the present-day Earth (1018 g). One
circularly polarized synchrotron radiation from amino acid, a-amino isobutyric acid, has been
a neutron star remnant of a supernova (Bonner identified in Antarctic micrometeorites (Brinton
1991). et al. 1998; Matrajt et al. 2004). These grains also
Strong infrared circular polarization, resulting contain a high proportion of metallic sulfides,
from dust scattering in reflection nebulae in the oxides, and clay minerals, a rich variety of inor-
Orion OMC-1 star-formation region, has been ganic catalysts which could have promoted the
observed (Bailey et al. 1998). Circular polariza- reactions of the carbonaceous material which
tion at shorter wavelengths might have been lead to the origin of life. Analysis of the dust
important in inducing chiral asymmetry in inter- grains collected by the Cosmic Dust mission sup-
stellar organic molecules that could be subse- ports a cometary origin for the micrometeorites
quently delivered to the early Earth (Bailey collected in Antarctica.
2001). A collection of CONCORDIA Antarctic
The discovery of a large number of meteorites micrometeorites recovered from ultraclean snow
since 1969 has provided new opportunities to close to Dome C provided the most unbiased
search for organic compounds in CM-type carbo- collection of large cosmic dust available. Many
naceous chondrites (Pizzarello et al. 2001; Glavin similarities can be found between Antarctic
et al. 2006; Pizzarello and Shock 2010) as well as micrometeorites and Wild 2 samples, in terms
in Renazzo-type (CR) chondrites found in Ant- of chemical, mineralogical, and isotopic compo-
arctica with natal enantiomeric excesses of up to sitions and in the structure and composition of
60 % (Pizzarello et al. 2012). their carbonaceous matter (Dobrica et al. 2009).
Micrometeorites Laboratory and Space Experiments

Dust collections in the Greenland and Antarctic Supporting Extraterrestrial Delivery
ice sheets (Maurette 1998, 2006) show that the
Earth captures interplanetary dust as micromete- Synthesis
orites at a rate of about 20,000 tonnes per year. Ultraviolet irradiation of dust grains in the inter-
About 99 % of this mass is carried by microme- stellar medium results in the formation of com-
teorites in the 50 to 500-mm size range. This value plex organic molecules. The interstellar dust
is about 2,000 times higher than the most reliable particles are assumed to be composed of silicate
estimate of the meteorite flux, i.e., about 10 tonnes grains surrounded by ices of different molecules,
including carbon-containing molecules. Ices of circularly polarized light of a given helicity,

H2O, CO2, CO, CH3OH, and NH3 were deposited inducing a stereo-specific photochemistry
at 12 K under a pressure of 107 mbar and irra- (Modica et al. 2014).
diated with electromagnetic radiation representa-
tive of the interstellar medium. The solid layer Space Travel
that developed on the solid surface was analyzed To estimate whether different amino acids could
by enantioselective gas chromatography and survive a trip in space embedded in micromete-
mass spectrometry GC-MS. After the analytical orites, a suite of amino acids like those detected
steps of extraction, hydrolysis, and derivatiza- in the Murchison meteorite has been exposed to
tion, 16 amino acids, including 6 protein amino space conditions in Earth orbit on board the
E
acids, were identified in the simulated ice mantle unmanned Russian satellites FOTON 8 and
of interstellar dust particles (Muñoz Caro 11, free and associated with clay minerals. Free-
et al. 2002). These amino acid identifications exposed aspartic acid and glutamic acid were
confirmed the preliminary amino acid formation partially destroyed during exposure to solar
obtained by Mayo Greenberg (Briggs et al. 1992). UV. However, decomposition was prevented
The chiral amino acids were racemic. Parallel when the amino acids were embedded in clays
experiments performed with 13C-containing sub- (Barbier et al. 1998). Amino acids have also been
stitutes definitely excluded contamination by bio- subjected to solar radiation outside the MIR sta-
logical amino acids. The results strongly suggest tion for 97 days. The samples were exposed in
that amino acids are readily formed in interstellar free form and associated with different ground
space. Irradiation experiments run with H2O, mineral supports to mimic micrometeorites, i.e.,
NH3, CH3OH, and HCN produced only three montmorillonite clay, powdered basalt, and pow-
amino acids, namely, glycine, alanine, and serine dered Allende meteorite. In the absence of min-
(Bernstein et al. 2002). eral protection, about half of the amino acids
Vacuum UV photo-irradiation of simpler ices were destroyed by UV radiation. The main pho-
containing H2O, CH3OH, and NH3 led to the tochemical degradation process was decarboxyl-
production of hydantoin, 2,4-imidazolidinedione ation. Significant protection from solar radiation
(De Marcellus et al. 2011), a molecule suspected was observed when the thickness of the added
to play an important role in the formation of minerals was 4–5 mm or greater (Boillot
oligopeptides. Hydantoin is known to form et al. 2002).
under extraterrestrial conditions, since it has The protective action of meteorite powder was
been detected in the soluble organic fraction of confirmed with the 18-month exposure experi-
primitive carbonaceous meteorites. Glycine and ment PROCESS using the EXPOSE-E facility
alanine were dominantly formed when mounted on the European Technology Exposure
5-substituted hydantoins were UV-irradiated Facility (EuTEF) platform on board the Interna-
and hydrolyzed (Sarker et al. 2013). tional Space Station. The amino acids exposed in
Irradiation of ice mixtures of H2O/13CH3OH/ the free form were degraded by more than 40 %,
NH3 by circularly polarized light provided while more than 80 % of the compounds associ-
16 distinct amino acids. The L-enantiomeric ated with meteorite powder were preserved
excesses (eeLs) were measured in five of (Cottin et al. 2012). The protective effect of the
them: a-alanine, 2,3-diaminopropionic acid, meteorite powder was confirmed with the 24
2-aminobutyric acid, valine, and norvaline, with month exposure in the EXPOSE-R facility,
values ranging from eeL = 0.20 % 0.14 % onboard the Russian module of the International
to eeL = 2.54 % 0.28 %. The results sup- Space Station (Bertrand et al. 2015).
port an astrophysical scenario in which the solar
system was formed in a high-mass star-forming Impact Shock Chemistry
region where icy grains were irradiated during the Intense bombardment probably caused some
protoplanetary phase by an external source of chemical reprocessing of the Earth’s primitive
atmosphere by impact shock chemistry. An indi- carbon, iron, nickel, water, and nitrogen to high-
cation of the number and timing of the impacts velocity impacts in a propellant gun. This exper-
onto the early Earth can be obtained by compar- iment simulated the chemistry experienced by
ison with the cratering record of the Moon, ordinary chondrites when hitting the Earth’s
which records impacts from the earliest history early oceans. They recovered several organic
of the solar system (Ryder 2003). Because of the molecules after the impact, including complex
larger size of the Earth and its greater gravita- molecules such as fatty acids and amines. Gly-
tional pull, about 20 times as many impacts cine, the simplest protein-building amino acid,
would have occurred on the early Earth as on was formed when the starting material contained
the Moon. Computer modeling of the impact ammonia, which is believed to have been formed
shock chemistry shows that the nature of the in prior impacts on the early Earth. To avoid
atmosphere strongly influences the shock prod- potential contamination by biological materials,
ucts (Fegley et al. 1986). A neutral CO2-rich the authors used solid carbon composed only of
13
atmosphere produces CO, O2, H2, and NO, C. Subsequent analyses of the recovered
while a reducing CO-rich atmosphere yields pri- organic products showed that they were all
marily CO2, H2, CH4, HCN, NH3, and formed with the 13C, ruling out the presence of
H2CO. The last three compounds are particu- any naturally formed biological contaminants,
larly interesting for prebiotic chemistry since which would have been enriched in 12C.
they can lead to amino acids via the Strecker The effects of impact shock on amino acids
synthesis. However, a CO-rich primitive atmo- and a peptide in artificial meteorites composed of
sphere may be unlikely. In laboratory experi- saponite clay were investigated (Bertrand et al.
ments, a gas mixture of methane, ammonia, 2009). The samples were subjected to pressures
and water subjected to shock heating followed ranging from 12 to 28.9 GPa, which simulated
by a rapid thermal quenching yielded the amino impact velocities of 2.4–5.8 km/s. Volatilization
acids glycine, alanine, valine, and leucine was determined by weight loss measurement, and
(Bar-Nun et al. 1970). Here again, the gas mix- the amino acid and peptide response was ana-
ture used does not represent a realistic primitive lyzed by gas chromatography-mass spectrome-
atmosphere, which was likely dominated by try. At the highest shock pressures, amino acids
CO2. Laboratory simulations of shocks were with an alkyl side chain were more resistant than
also run with a high-energy laser. CH4- those with functional side chains. Impact shock
containing mixtures generated hydrogen cya- may therefore act as a selective filter to
nide and acetylene, but no organics could be the delivery of extraterrestrial amino acids via
obtained with CO2-rich mixtures (McKay and carbonaceous chondrites. The peptide cleaved
Borucki 1997). into its two primary amino acids. Some chiral
Large impacts could also have promoted some amino acids experienced partial racemization
fine ejecta particle chemistry. For 10–15-km- during the course of the experiment, suggesting
sized impactors, modeling (Lyons and Vasavada that the enantiomeric excesses measured in
1999) has shown that 100-mm fine ejecta particles carbonaceous chondrites are probably under
would have been heated to about 200 C during evaluated.
reentry in a 0.3 bar CO2 atm. For impactors larger
than 20 km, the heat experienced by the fines Conclusion
would destroy all organics by pyrolysis or Meteorite and micrometeorite collection and
combustion. analysis as well as laboratory and space experi-
Furukawa and colleagues (Furukawa ments strongly support an extraterrestrial path-
et al. 2009) investigated whether, rather than way for the delivery of organic molecules at the
just transporting amino acids, meteorite impacts time that life originated. The building blocks of
could themselves synthesize organic com- life or their precursors could have been brought in
pounds. The group subjected a mixture of solid by small impactors, possibly supplemented by
prebiotic molecules formed in the atmosphere Cottin H, Guan YY, Noblet A, Poch O, Saiagh K, Cloix M,
and oceans by meteorite impacts. Macari F, Jérome M, Coll P, Raulin F, Stalport F,
Szopa C, Bertrand M, Chabin A, Westall F,
Chaput D, Demets R, Brack A (2012) The PROCESS
experiment: an astrochemistry laboratory for solid and
See Also gaseous organic samples in low-earth orbit. Astrobiol-
ogy 12:412–425
▶ Comet Cronin JR, Pizzarello S (1997) Enantiomeric excesses in
meteoritic amino acids. Science 275:951–955
▶ Meteorites De Marcellus P, Bertrand M, Nuevo M, Frances Westall F,
Le Sergeant d’Hendecourt L (2011) Prebiotic signifi-
cance of extraterrestrial ice photochemistry: detection
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Maurette M (2006) Micrometeorites and the mysteries of

our origins. Springer, Berlin Extreme Environment
Maurette M, Brack A (2006) Cometary petroleum in
Hadean time? Meteorit Planet Sci 41:5247
McKay CP, Borucki WJ (1997) Organic synthesis in Felipe Gomez
experimental impact shocks. Science 276:390–392 Centro de Astrobiologı́a (CSIC/INTA), Instituto
Modica P, Meinert C, de Marcellus P, Nahon L, Nacional de Técnica Aeroespacial, Torrejón de
Meierhenrich UJ, Le Sergeant d’Hendecourt L
(2014) Enantiomeric excesses induced in amino acids Ardoz, Madrid, Spain
by ultraviolet circularly polarized light irradiation of
extraterrestrial ice analogs : a possible source of asym-
metry for prebiotic chemistry. Astrophys J 788:79 Keywords
Muñoz Caro GM, Meierhenrich UJ, Schutte WA,
Barbier B, Arcones Segovia A, Rosenbauer H,
Thiemann WH-P, Brack A, Greenberg JM (2002) Extremophiles
Amino acids from ultraviolet irradiation of interstellar
ice analogues. Nature 416:403–405
Pizzarello S (2007) The chemistry that preceded life’s
origin: a study guide from meteorites. Chem Biodivers Synonyms
4:680–693
Pizzarello S, Huang Y (2005) The deuterium enrichment Extreme field sites
of individual amino acids in carbonaceous meteorites:
a case for the presolar distribution of biomolecules
precursors. Geochim Cosmochim Acta 69:599–605
Pizzarello S, Shock E (2010) The organic composition of Definition
carbonaceous meteorites: the evolutionary story ahead
of biochemistry. Cold Spring Harb Perspect Biol 2:
a002105 An extreme ▶ environment is a habitat charac-
Pizzarello S, Huang Y, Becker L, Poreda RJ, Nieman RA, terized by harsh environmental conditions,
Cooper G, Williams M (2001) The organic content of beyond the optimal range for the development
the Tagish Lake meteorite. Science 293:2236–2239
of humans, for example, pH 2 or 11, 20 C or
Pizzarello S, Zolensky M, Turk KA (2003) Non racemic
isovaline in the Murchison meteorite: chiral distribu- 113 C, saturating salt concentrations, high radi-
tion and mineral association. Geochim Cosmochim ation, and 200 bars of pressure, among others.
Acta 67:1589–1595 Basically, these are all inhospitable conditions
Pizzarello S, Schrader DL, Monroe AA, Lauretta DS
for life. By definition, the organisms that are
(2012) Large enantiomeric excesses in primitive mete-
orites and the diverse effects of water in cosmochemical able to live in extreme environments are
evolution. Proc Natl Acad Sci U S A 109:11949–11954 known as ▶ extremophiles. Not so long ago it
Ryder G (2003) Bombardment of the Hadean Earth: was thought that life could not occur under
wholesome or deleterious? Astrobiology 3:3–6
extreme conditions. In the 1960s, Professor
Sarker PK, Takahashi J-I, Obayashi Y, Kaneko T,
Kobayashi K (2013) Photo-alteration of hydantoins Thomas D. Brock, from Wisconsin-Madison
against UV light and its relevance to prebiotic chem- University, isolated and described the first
istry. Adv Space Res 51:2235–2240 organisms from Yellowstone National Park,
Schmitt-Kopplin P, Gabelica Z, Gougeon RD, Fekete A,
USA. This organism, Thermus aquaticus, is
Kanawati B, Harir M, Gebefuegi I, Eckel G, Hertkorn
N (2010) High molecular diversity of extraterrestrial capable of growing at temperatures higher than
organic matter in Murchison meteorite revealed 70 C. Its DNA polymerase has been widely
40 years after its fall. Proc Natl Acad Sci U S A applied in molecular biology as it is the base of
107:2763–2768
the polymerase chain reaction (PCR) based on
Stoks PG, Schwartz AW (1982) Basic nitrogen-
heterocyclic compounds in the Murchison meteorite. the thermophilic properties of the microorgan-
Geochim Cosmochim Acta 46:309–315 ism that produces it. Sulfolobus acidocaldarius,
Yabuta H, William LB, Cody GD, Alexander CMO’D, growing at temperatures higher than 85 C, was
Pizzarello S (2007) The insoluble carbonaceous mate-
also isolated and characterized at the same time
rial of CM chondrites: a possible source of discrete
compounds under hydrothermal conditions. Meteorit and was the first characterized hyperthermo-
Planet Sci 42:37–48 philic archaea.
Extreme Environment 829
History 1. Organisms developing in these acidic envi-

ronments are known as acidophiles.
In 1974, R. D. MacElroy published “Some com- Extreme alkaline environments are those
ments on the evolution of extremophiles” with a pH above 9. Examples of this type of
(Biosystems 6: 74–75); this was the first time environment are the soda lake of Magadi,
the term “▶ extremophile” was used. Mono Lake, or saltpans. Organisms found in
these environments are called alkaliphiles.
Extreme ionic strength. Hypersaline environ-
Overview ments have an ionic concentration higher
than of seawater, >3.5 %. Typical hypersaline
E
Depending on the extreme physicochemical con- environments are the Dead Sea (Israel), the
ditions that characterize the extreme environ- Great Salt Lake (USA), or the salterns of
ments, they are classified as follows: Santa Pola (Spain). Organisms able to grow
at high ionic strength are known as halophiles.
Extreme temperature. Two types of extreme eco- Most alkaliphilic organisms are also
systems can be described: cold and hot. halophiles.
Extremely cold environments are those with Extreme pressure environments are those under
temperatures consistently below 5 C. They extreme hydrostatic or litho pressure, such as
can be found in deep ocean niches, at the aquatic habitats at depths of 2,000 m or more
peaks of high mountains, or in the polar or deep-subsurface ecosystems. Organisms
regions. Organisms living in extreme cold living under high pressure can be classified
environments are known as psychrophiles. as barotolerant if they can tolerate high pres-
Some of them, such as the organisms found sure or barophilic if they depend on pressure
in Vostok Lake are able to live at 20 C. to grow.
Extremely hot environments are character-
ized by regular temperatures higher than High-radiation environments are those habi-
45 C. These environments are typically tats exposed to abnormally high radiation doses,
influenced by geothermal activity as geysers including ultraviolet or infrared radiation, like
and fumaroles of continental volcanic areas or deserts, the top of high mountains, or on the
deep-sea vents. Typical extreme hot environ- surface of the International Space Station.
ments can be found in the geothermal areas of Xeric environments are extreme dry habitats
Yellowstone and in some locations of Iceland with seriously limited water, an extremely impor-
or Kamchatka. The organisms in these envi- tant element for life. Cold and hot deserts are
ronments are thermophiles. Some microor- some examples of these extreme environments.
ganisms are able to develop at temperatures Oligotrophic environments are extreme eco-
higher than 80 C, and they are called systems that offer low levels of nutrients to sus-
hyperthermophiles. These organisms are asso- tain life. Oligotrophic environments include
ciated with hydrothermal activities. deep oceanic sediments, polar ice, or the deep
Extreme pH. Extreme environments can be clas- subsurface.
sified as acidic or alkaline according to
their pH.
Extreme acidic environments are natural See Also
habitats in which the pH is below 5. Some
examples of extreme acidic environments are ▶ Acidophile
Rı́o Tinto (Iberian Pyritic Belt, SW Spain), ▶ Alkaliphile
mean pH 2.3, or Iron Mountain in California ▶ Compatible Solute
(USA) where the pH in some areas is below ▶ Deep-Sea Microbiology
830 Extreme Field Sites
▶ Deep Subsurface Microbiology Definition

▶ Desiccation
▶ Endolithic Extreme Ultraviolet light, EUV, corresponds to
▶ Environment photons with wavelengths between 100- and
▶ Extremophiles 10 nm (12.4–124 eV).
▶ Halophile Ultraviolet Radiation
▶ Hyperthermophile
▶ Piezophile
▶ Psychrophile See Also
▶ Solar UV Radiation, Biological Effects ▶ VUV
▶ Terrestrial Analog ▶ Wavelength
Amaral-Zettler L, Gómez F, Zettler E, Keenan BG,

Amils R, Sogin M (2002) Eukaryotic diversity in Extremophiles
Spain’s river of fire. Nature 417:137. doi:10.1038/
417137a Daniel Prieur
Karl DM, Bird DF, Björkman K, Houlihan T, Shackelford R,
Tupas L (1999) Microorganisms in the Accreted Ice of Université de Bretagne Occidentale (University
Lake Vostok, Antarctica. Science 286(5447):2144–2147. of Western Britanny), Brest, France
doi:10.1126/science.286.5447.2144 Institut Universitaire Européen de la Mer
Rothschild LJ, Mancinelli RL (2001) Life in extreme (IUEM), Technopôle Brest–Iroise, Plouzané,
environments. Nature 409:1092–1101. doi:10.1038/
35059215
France
Wharton DA (2002) Life at the limits: organisms in
extreme environments. Cambridge University Press,
Cambridge. ISBN 0521782120 Keywords
Acidophile; Alkaliphile; Extremophile; Halo-

phile; Hydrostatic pressure; pH; Piezophile;
Extreme Field Sites Psychrophile; Salt concentration; Temperature;
Thermophile
Synonyms
Extremophilic organisms; Life, limits of

Extreme Ultraviolet Light
José Cernicharo Quintanilla Definition

Department of Astrophysics, Laboratory of
Molecular Astrophysics, Iorrejón de Ardoz, Life is influenced by physical parameters such
Madrid, Spain as temperature, pH, salinity, pressure,
etc. Extremophiles are organisms that thrive in
ecosystems where at least one physical parameter
Synonyms is close to the known limits of life with respect to
this parameter. While some organisms may tem-
EUV porarily survive harsh conditions by forming
Extremophiles 831
resistant stages (spores) or through specific (millions of years in the view of some authors)
mechanisms (heavy metal resistance), true and then produce growing cells after environ-
extremophiles require these conditions. For mental conditions become favorable. Other
instance, hyperthermophiles successfully com- organisms can temporarily resist toxic com-
plete their life cycle at optimal temperatures pounds such as heavy metals or ionizing radia-
above 80 C and commonly do not grow at all tion, through specific mechanisms. But these
below 60–70 C. two categories do not require such hostile con-
ditions for growth. A third category truly
deserves the name extremophile: These organ-
History isms require environmental conditions hostile
E
for most of the other organisms to complete
Before 1965, the upper temperature limit known their whole life cycle: very low temperatures
for life was about 73 C. When Thomas Brock for psychrophiles, very high temperature for
discovered thermophilic bacterium Thermus thermophiles, very low pH for acidophiles,
aquaticus in the Octopus Hot Spring in Yellow- very high pH for alkaliphiles, elevated salt con-
stone National Park (USA), the research on centrations for halophiles, or elevated hydro-
extremophiles truly began. Thermus aquaticus static pressure for piezophiles.
lives in temperatures ranging from 50 C to
80 C. Since then, many alkaliphiles, acidophiles,
halophiles, and piezophiles have been discovered Basic Methodology
and studied for their taxonomy, ▶ phylogeny,
physiology, biochemistry, molecular biology, Since extremophiles thrive in environments hos-
genetics, and applications. According to Gerday tile for many organisms and particularly human
and Glansdorff (2007), “For biologists, the reali- beings, the main difficulty encountered during
zation that many forms of life are actually con- the study of extremophiles is access to extreme
fined to environments that are severely hostile by environments. Whereas access to continental hot
human standards will remain one of the most springs (acidic or alkaline), glaciers, salt mines,
significant achievements of the second half of or evaporating ponds only requires specific
the twentieth century.” But “the questions raised clothing and caution, access to the deep-sea
by the molecular basis and the emergence of hydrothermal vents, where psychro-piezophiles
different forms of extremophily are not only or hyperthermophiles are found, is much more
deep and varied, but, not surprisingly, many complex. In this case, sophisticated underwater
remain controversial.” At the beginning of the vehicles (manned submersibles or remotely
twenty-first century, research on extremophiles operated vehicles (ROV)) are needed. The next
is still an open field. challenge after sampling is the preservation of
samples for further cultivation in the laboratory.
Many extremophiles are strictly anaerobic and
Overview require anaerobic-reduced conditions during
preservation at low temperature. Probably
Living organisms exposed to severe environ- many obligate piezophiles do not survive sam-
mental conditions are frequently named pling procedures since it is very difficult (and
“extremophiles.” However, they belong to dif- very expensive) to deploy pressure-retaining
ferent categories. Some organisms resist hostile samplers. Psychrophiles are very sensitive to
conditions (elevated temperatures, ▶ desicca- moderate temperature (room temperature), and
tion, etc.) by forming sophisticated resistant all the glassware and reagents used must be
and dormant stages such as the spores produced precooled.
by several Gram-positive ▶ bacteria. Such Finally, cultivation of extremophiles requires
forms can survive for very long periods mimicking the main environmental conditions
832 Extremophiles
of their environments in the laboratory. The diffi- never grow beyond 20 C. Their minimum tem-
culties of growing lithoautotrophic organisms perature for growth can be below 0 C. Organisms
even under non-extreme conditions are great, but that can grow at low temperatures but also beyond
the most difficult to grow are probably piezophiles 20 C are named psychrotolerant. The lower tem-
which require high-pressure bioreactors. perature limit for growth is not easy to determine,
since very low temperatures (80 C in deep
freezer, 196 C in liquid nitrogen) are used for
Key Research Findings long-term preservation of both prokaryotic and
eukaryotic cells. However, growth of bacterial
Thermophiles cells at temperatures as low as 10 to 12 C,
Thermophiles are organisms that grow optimally and metabolic activity at 20 C in concentrated
above 60 C, and hyperthermophiles grow opti- brines have been reported. Psychrophiles not
mally above 80 C. The most thermophilic and only inhabit the deep cold oceans but also
hyperthermophilic organisms are prokaryotes polar regions, high mountains, glaciers, etc.
and belong to the Bacteria, but essentially They almost certainly belong to many phyloge-
▶ Archaea domains. They have been isolated netic lineages, but most of the cultivated species
from diverse hot environments such as continen- belong to the Bacteria domain, particularly the
tal hot springs, oil reservoirs, or deep-sea hydro- proteobacteria.
thermal vents. According to their natural
environments, many of them combine Halophiles
thermophily with acidophily or piezophily. Halophiles are organisms living in salty environ-
They have representatives in various metabolic ments such as the oceans. However, many envi-
types: They may be aerobes, microaerophiles or ronments are much more saline and salt
anaerobes, and chemoorganotrophs or concentrations can reach 340 g/l. Typical settings
chemolithoautotrophs. As a whole, they may are salt lakes, solar salterns, underground deposits
use a variety of electron donors and acceptors. of rock salts, but also salted food products. Organ-
Many of them have very short generation times, isms living optimally in these highly salted envi-
about 30 min under optimal conditions. ronments are named extreme halophiles. Several
The most thermophilic organism is an salty environments also have a high pH and a
Archaea, Pyrolobus fumarii, isolated from a rather high temperature, and consequently their
deep-sea hydrothermal vent. P. fumarii grows in inhabitants are polyextremophilic. Extreme halo-
a temperature range of 90–113 C, with an opti- philes (growing in salt concentrations above 100 g/
mum at 106 C. It can survive an exposure of 24 h l) are found in the three domains of life, but most of
in an autoclave. Growth at 121 C of an iron them are prokaryotes, and the most extreme belong
reducer organism, also isolated from a deep-sea to the Archaea domain. Two lineages of
hydrothermal vent, has been reported. However, halobacteriales and certain methanogens are
the strain has not been published as a novel included. The halobacteriales are heterotrophic
organism nor deposited in type culture collec- organisms that produce red pigments, some of
tions, and this result is still to be confirmed. which (bacteriorhodopsin, halorhodopsin) are
Thermophiles may also harbor viruses that live involved in light-driven proton and chloride
in the same environmental conditions, but their pumps.
life cycle and relationships with their hosts are
still unknown. Acidophiles
Acidophilic organisms grow optimally at pH
Psychrophiles below 7, but there is a distinction between mod-
Psychrophiles are cold-adapted organisms that erate acidophiles that grow at pH from 5 to 3 and
grow optimally at temperatures around 15 C and extreme acidophiles that grow at pH below
Extremophiles 833
3. Acidophiles are found in natural acidic isolated from a dead crustacean amphipod col-
springs in volcanic areas, but also in sulfide ore lected at the Mariana Trench (10,470 m depth). It
or coal deposits. Acidophiles exist in the three is an aerobic heterotrophic bacterium that grows
domains of life, and several yeasts, fungi, and optimally at 2 C (it is also a ▶ psychrophile),
protozoa grow at pH below 2. Among the pro- under 690 bars (or 69 Mpa) with a generation
karyotes, acidophiles are very diversified from a time of 25 h. When exposed to hydrostatic pres-
metabolic point of view, and both heterotrophs sure, this organism loses its ability to form colo-
and autotrophs are found. Many acidophiles oxi- nies and dies. Piezophilic characteristics are also
dize sulfides and sulfur, using oxygen as an elec- known in some hyperthermophiles isolated from
tron acceptor. Since ferrous iron is stable at low deep-sea hydrothermal vents. Pyrococcus
E
pH under aerobic conditions, many acidophiles yayanosii strain CH1 grows in a temperature
are also iron oxidizers. The most acidophilic range of 80–105 C, with an optimum at 98 C,
organism on Earth is the Archaea Picrophilus, under an optimal hydrostatic pressure of 52 Mpa.
which has an optimal pH of 0.7 and still grows at Its pressure range extends from 20 to at least
pH 0. 120 Mpa. At all temperatures, no growth was
observed for pressures below 20 Mpa.
Alkaliphiles
Alkaliphiles are commonly divided into two cat-
egories: the alkali-tolerant organisms can grow Applications
at alkaline pH (8–9), but their optimum is around
neutral pH. True alkaliphiles can grow at pH Extremophiles are able to grow under extreme
above 10 and/or grow equally well or better conditions of temperature, pH, salt concentration,
(rate, yield) at pH above 9. Some are facultative and pressure. Consequently, the metabolic path-
▶ alkaliphile; true alkaliphiles cannot grow at ways and enzymes involved in these pathways
pH below 8. Alkaliphiles occur in natural high- are also extremophilic and are named
pH environments such as ▶ soda lakes, under- extremozymes. Extremozymes have two general
ground alkaline waters, and hidden small niches properties: they are active under extreme condi-
such as insect guts. They are also found in arti- tions, but are also very stable. For those reasons
ficial environments such as waste of food- they represent a powerful tool for industrial bio-
processing industries. They have representatives transformations, carried out under harsh condi-
in the three domains of life, but are most abun- tions. Most of them come from thermophilic and
dant in several lineages of the Bacteria domain, hyperthermophilic organisms.
particularly the low G + C Gram-positive bacte- Many extremozymes are involved in starch
ria of the genus Bacillus. Some alkaliphiles also processing: alpha- and beta-amylases,
exhibit slight thermophilic properties. glucoamylases, alpha-glucosidase, pullulanase,
CgTases, branching enzymes, and amylomaltases.
Piezophiles There are extremozymes that can be used for other
Piezophiles were first named barophiles, from the transformations such as cellulose, xylan, chitin,
Greek word for weight. Piezophile is more appro- pectin-degrading enzymes, proteolytic enzymes,
priate because it comes from the Greek word for lipases and esterases, alcohol dehydrogenases,
pressure. Piezophiles thrive in environments glucose and arabinose isomerases, nitrile-
exposed to elevated hydrostatic pressure. degrading enzymes, etc. Modern DNA technology
Piezotolerant, piezophilic, and obligate also utilizes extremozymes from thermophiles.
piezophilic exist. The latter requires pressures The most famous example is the thermostable
above 1 bar to grow. They are found in habitats Taq polymerase, from the thermophilic bacterium
such as deep oceans, deep aquifers, oil fields, and Thermus aquaticus. This polymerase permitted
deep sediments. The first obligate piezophile was the revolution in DNS technology through the
834 Extremophiles
PCR (polymerase chain reaction) technology. By pushing the definition of the limits for life
Other enzymes such as ligases, nucleases, and to ever greater extremes, biologists engaged in
topoisomerases are also commonly used. research on extremophiles will help develop
Extremophiles can also be used as whole-cell novel hypotheses about the origin of life on
biocatalysts for biomining, decontamination or Earth and its existence elsewhere.
bioremediation, and hydrogen production.
Finally, biomolecules extracted from
extremophiles such as proteins and peptides, bio- See Also
polymers, compatible solutes, or lipids have a
wide range of applications. ▶ Acidophile
▶ Alkaliphile
▶ Anaerobe
Future Directions ▶ Archaea
▶ Bacteria
At the beginning of the twenty-first century, the ▶ Chemolithoautotroph
world of extremophiles has revealed a variety of ▶ Deep-Sea Microbiology
fascinating organisms, most of which are pro- ▶ Deep Subsurface Microbiology
karyotes. However, despite the thousands of ▶ Desiccation
papers that have been published on the subject, ▶ Ecosystem
many questions remain. ▶ Enzyme
Extremophiles thrive at the limits of the life on ▶ Eukarya
Earth, but what those limits are precisely is not ▶ Extreme Environment
yet fully understood, particularly for elevated ▶ Halophile
temperatures and pressures. Although it is not ▶ Hot Spring Microbiology
possible to imagine an organism more acidophilic ▶ Hot Vent Microbiology
than Picrophilus (already at the low pH limit), it ▶ Hydrothermal Vent Origin of Life Models
may be possible to find an organism that com- ▶ Hyperthermophile
bines extreme acidophily with extreme ▶ Lithotroph
thermophily and piezophily or any other combi- ▶ PCR
nation. To discover such pluri-extremophiles, ▶ Piezophile
exploration of planet Earth for new biotopes ▶ Prokaryote
must be continued. It was recently reported that ▶ Psychrophile
living microorganisms exist in very deep ▶ Soda Lakes
(1,600 m thickness) and old (110 million years) ▶ Thermophile
marine sediments. At this location, temperature
was estimated to be only 80–100 C. But the References and Further Reading
known temperature limit for life is 113 C
(perhaps more), and because temperature Corliss JB et al (1979) Submarine thermal springs on the
increases by only 30 C per kilometer, life could Galapagos rift. Science 203:1073–1083
occur at even greater depths. Gerday C, Glansdorff N (2007) Physiology and biochem-
istry of extremophiles. ASM, Washington, DC
Such organisms may also represent novel lin- Kato C (1999) Barophiles (piezophiles). In: Horikoshi K,
eages and may use novel metabolic pathways or Tsujii K (eds) Extremophiles in deep-sea environ-
novel enzymes. For each of them, the cellular and ments. Springer, Tokyo, pp 91–111
Lopez-Garcia P (2005) Extremophiles. In: Gargaud M,
molecular mechanisms of adaptation remain to
Barbier B, Martin H, Reisse J (eds) Lectures in astro-
be understood and may open a variety of potential biology. Advances in astrobiology and biogeophysics,
applications. vol II. Springer, Berlin, pp 657–682
Extremophilic Organisms 835
Prieur D, Marteinsson VT (1998) Prokaryotes living under Zeng X, Birrien JL, Fouquet Y, Cherkashov G,
elevated hydrostatic pressure. Adv Biochem Eng Jebbar M, Querellou J, Oger P, Cambon-Bonavita
Biotechnol 61:23–35 MA, Xiao X, Prieur D (2009) Pyrococcus CH1, an
Prieur D et al (2010) Piezophilic prokaryotes. In: Sebert obligate piezophilic hyperthermophile: extending the
P (ed) Comparative high pressure biology. Science upper pressure-temperature limits for life. ISME
Publishers, Enfield (NH), Jersey, pp 285–322 J 3:873
Roussel E et al (2008) Extending the sub sea-floor bio-
sphere. Science 320:1046
Xiang Z et al (2009) Pyrococcus CH1, an obligate
piezophilic hyperthermophile isolated from a deep-
sea hydrothermal vent. ISME J 1–4
Yayanos AA (1986) Evolutional and ecological implica-
Extremophilic Organisms
tions of the properties of deep-sea barophilic bacteria. E
Proc Natl Acad USA 83:9542–9546 ▶ Extremophiles
F
Facula, Faculae ▶ Impact Basin

▶ Jupiter
Roland J. Wagner ▶ Macula, Maculae
German Aerospace Center (DLR), Institute of ▶ Satellite or Moon
Planetary Research, Berlin, Germany ▶ Saturn
▶ Titan
Definition
A facula is a bright area on the icy satellites of

Faint Young Sun Paradox
▶ Jupiter, ▶ Ganymede, ▶ Callisto, and
Amalthea and on ▶ Saturn’s satellite ▶ Titan.
Manuel G€udel
Faculae on Ganymede and Callisto are circular
Department of Astrophysics, University of
or elliptical and up to several hundred kilometers
Vienna, Vienna, Austria
in diameter. Faculae are thought to have been
created by impacts into the icy ▶ crusts of these
two Jovian satellites, possibly with plastic or
Keywords
liquid material present in the subsurface. Titan
shows two globally abundant surface units char-
Climate; Greenhouse gases; Solar mass loss;
acterized by either bright or dark ▶ albedo. Fac-
Young Sun
ulae on this satellite are irregularly shaped,
represent slivers or islands of bright terrain, are
located within extensive areas of dark terrain, and
are possibly of nonimpact origin. Synonyms
Faint young Sun problem

See Also
▶ Albedo Feature Definition

▶ Callisto
▶ Crater, Impact The apparent contradiction between model cal-
▶ Crust culations that indicate a dimmer young Sun and
▶ Ganymede consequently mean below-freezing temperatures
DOI 10.1007/978-3-662-44185-5
838 Faint Young Sun Paradox
at the Earth’s surface and the geological evidence atmosphere follows from the energy balance
that early climates on Earth and Mars were mild if between optical/near-infrared emission irradiat-
not hot and that liquid water was abundantly ing the planet and mid-infrared thermal radiation
present on the other hand. that is lost to space. The incoming power from the
Sun is
Overview Pin ¼ pR2 Sð1 AÞ
There is clear evidence for a mild climate on the where S = LJ/4pd2 = 1,366 W m2 is the solar
young Earth and Mars, allowing liquid water – a constant (solar radiative energy flux at the dis-
prerequisite for the formation of life as we know tance d of the Earth, where LJ is the solar lumi-
it – to exist on the surface of both planets at ages of nosity), A 0.29 is the Earth’s Bond albedo
a few 100 Myr to 1 Gyr. Liquid water seems to have (fraction of radiation that is reflected), and
subsequently disappeared from the surface of Mars. R = 6,378 km is the Earth’s radius. The power
However, detailed stellar evolutionary theory leaving from the Earth’s atmosphere can be
applied to the Sun during its main-sequence life approximated as radiation from a blackbody sur-
indicates that it was fainter by about 30 % when it face with temperature T,
arrived on the main sequence 4.6 billion years ago,
and the radiative output increased only slowly in Pout ¼ 4pR2 esT 4
the next billion years. Both planetary surfaces
would thus have been completely frozen. To solve where s = 5.67 108 W m2 K4 is the
this apparent paradox (Sagan and Mullen 1972), Stefan-Boltzmann constant and e is the surface
various hypotheses have been proposed although a mid-infrared emissivity. Equating Pin = Pout
conclusive answer is still outstanding. The most leads to
popular theory assumes higher admixtures of
greenhouse gases in the atmospheres of Earth and Sð 1 AÞ
T4 ¼
Mars; such gases include carbon monoxide, ammo- 4es
nia, and methane (CO2, NH3, and CH4, respec-
tively). A radically different hypothesis posits that With the above numbers and e 0.9 for solid
the young Sun was not faint – it may rather have rock (Sagan and Mullen 1972), T 260 K. For
been brighter because it was more massive by a few a planet surrounded by an atmosphere, the effec-
percent. Direct evidence for the implied strong tive temperature can be raised owing to the pres-
mass loss in the younger epochs of solar evolution ence of greenhouse gases. The latter are nearly
is still outstanding. Further hypotheses assume a transparent to incident optical and near-infrared
lower albedo due to less efficient cloud formation, light but strongly absorb reemitted thermal radi-
for example, as a consequence of more efficient ation in the mid-infrared. The equation for the
suppression of cosmic rays in the young solar sys- mean surface temperature of the atmosphere can
tem, or a smaller fractional area of land, or the lack then be approximated by (e.g., Feulner 2012)
of biologically induced cloud condensation nuclei.
For a recent review of the subject and a summary of Sð 1 AÞ 3t
T ¼
4
1þ
evidence for a warm early Earth with liquid surface 4es 4
water, see Feulner (2012).
where t* is the column infrared gray opacity
responsible for the warming by greenhouse gases.
Basic Methodology Heat flow from the interior of the Earth is, in
contrast, negligibly small (although it matters for
The average, effective equilibrium temperature maintaining plate tectonics; Feulner 2012 and
of a planetary surface in the absence of an references therein). Considering the greenhouse
Faint Young Sun Paradox 839
effect for a present-day atmosphere leads to a pieces of evidence indicate mild climates and
somewhat elevated average temperature of the presence of liquid water on both young
288 K (15 C), in agreement with measurements planets (Valley et al. 2002; Feulner 2012).
(e.g., Sagan and Mullen 1972; Kasting and
Catling 2003).
The total radiative output of the young Sun can Key Research Findings
be assessed from two sources: (1) The theory of
stellar evolution indicates that the young Sun at a Assuming the same atmospheric composition for
time when it started core hydrogen burning on the the Earth as today but a solar constant reduced by
main sequence emitted 30 % less electromagnetic about 30 %, calculations yield average atmo-
radiation than at present (e.g., Sackmann and spheric temperatures of about 260 K for the
Boothroyd 2003). (2) The total luminosity of young Earth around 4 billion years ago. The F
stars with known masses and ages can in principle temperature would have remained below the
be derived from observations; stellar masses and freezing point until 2 billion years ago (Sagan
ages are, however, difficult to infer accurately. and Mullen 1972; Kasting and Catling 2003;
Our knowledge of the early climate history of Fig. 1). This clearly contradicts geological evi-
the Earth and other planets, in particular Mars, dence for a significantly warmer early climate on
relies on geological evidence, such as (1) oxygen Earth (e.g., Kasting and Toon 1989) and espe-
isotopes measured in Jack Hills zircons dated at cially also on Mars. This contradiction is the
4.4–4.3 Ga, which suggested that parent rocks essence of the faint young Sun paradox (FYSP).
interacted with liquid water; (2) 3.8 Ga old sedi- Various hypotheses have been put forward
mentary rocks from Isua (West Greenland) addressing the FYSP but the solution remains
clearly deposited in aquatic environments; and inconclusive. We summarize the key points of
(3) extensive outflow channels and valley net- the proposed solutions below. Considering the
works in the Martian highlands. All of these above equation for the atmospheric
300 1.0
Solar luminosity relative
to pressent value
Freezing point of water

Temperature (K)
275 0.9
TS
S/S0
250 0.8
Te
225 0.7
4 3 2 1 0
Billions of years before present
Faint Young Sun Paradox, Fig. 1 Illustration of the affected by the greenhouse (with fixed CO2 mixing ratio
faint young Sun paradox for the Earth, in the context of the and relative humidity) (From Kasting and Catling 2003,
atmospheric greenhouse. The solid line depicts the solar reprinted, with permission, from the Annual Review of
luminosity relative to the present value (right y-axis); the Astronomy and Astrophysics, volume 41 # 2003 by
lower dashed curve is the effective temperature of the Annual Reviews, www.annualreviews.org. Courtesy of
Earth without atmosphere, and the upper dashed curve J. Kasting; credit: Scientific American and Princeton Uni-
shows the calculated, mean global surface temperature versity Press)
temperature, T, and the determining variables, Another leading candidate among greenhouse
most potential solutions of the FYSP for the gases is ammonia (NH3) which would be required
young Earth fall into the following categories: in only relatively small amounts. However, NH3
(i) an increased greenhouse effect (t ), (ii) a dissociates rapidly if subject to solar UV radia-
reduced albedo A, and (iii) an increased solar tion (Kuhn and Atreya 1979), but additional
luminosity Lo. A smaller semimajor axis of the methane (CH4) may produce a high-altitude
Earth’s orbit (resulting in an increased solar flux haze of organic solids through photolysis, which
S at Earth) or a significantly reduced emissivity e shields ammonia sufficiently from UV dissocia-
are unlikely (Feulner 2012 and references tion (Sagan and Chyba 1997). Methane itself is an
therein.) efficient greenhouse gas (Kasting 1997; Pavlov
et al. 2000). But the mixing ratio of CH4:CO2
Different Atmospheric Composition: should stay below unity to avoid increased albedo
Greenhouses due to haze formation and therefore an anti-
The composition of the young atmospheres may greenhouse effect (McKay et al. 1999; Pavlov
have been different, allowing for much stronger et al. 2001; Haqq-Misra et al. 2008), in turn
greenhouses on Earth or Mars. Many authors requiring too high levels of CO2 for a significant
have favored this hypothesis, although it faces greenhouse. Haze is produced at even consider-
its own problems. Larger amounts of CO2 ably lower CH4:CO2 mixing ratios (DeWitt
would strongly support the greenhouse together et al. 2009). On the other hand, enhanced levels
with water vapor (Kasting 1993). The required of H2 suppress haze formation and this limits the
CO2 level to keep liquid water oceans would be anti-greenhouse effect, possibly resulting in a net
self-regulated by the carbonate-silicate cycle greenhouse warming by CH4+CO2 (DeWitt
(e.g., Walker et al. 1981; Kasting and Catling et al. 2009). The presence of sufficiently high
2003). Increased CO2 levels would at the same levels of H2 remains to be demonstrated, how-
time also prevent massive losses of N2 under the ever. Ethane (C2H6) and carbonyl sulfide
influence of an enhanced early solar wind and (OCS) have been proposed as alternative efficient
enhanced extreme-ultraviolet radiation greenhouse gases in the young terrestrial
(Lichtenegger et al. 2010). There are, however, atmosphere (Haqq-Misra et al. 2008; Ueno
various geochemical and geological evidences et al. 2009) although OCS itself, for example,
against the required very massive CO2 atmo- is rapidly photodissociated (Domagal-Goldman
spheres, such as the absence of siderite in et al. 2011).
paleosols (Rye et al. 1995; Rosing et al. 2010,
and references therein); a dense Martian CO2 Albedo Effects Due to Cloud Formation and
atmosphere would significantly enhance the Mar- Land Coverage
tian albedo, thus not raising the temperature suf- A much lower albedo, A, would allow T to rise.
ficiently (Kasting 1991). Although these same This explanation was initially deemed unlikely
clouds may also backscatter thermal radiation because the probably larger surface of the young
and therefore support the greenhouse mechanism Earth covered by ice would rather increase
(Forget and Pierrehumbert 1997), experiments A (Sagan and Mullen 1972).
suggest that this effect is too small to raise the The albedo could, however, have been lower
temperatures above the freezing point (Glandorf due to a lower cloud coverage in particular in
et al. 2002). However, for a period around cooler environments (Rossow et al. 1982;
2–2.5 Gyr ago, recent calculations indicate that Charlson et al. 1987; Rosing et al. 2010). Detailed
a lower CO2 content of 2.9 Mb partial pressure studies of clouds show two competing effects
would have been sufficient to keep liquid water (Goldblatt and Zahnle 2011a): low-lying clouds
on the Earth’s surface, and this pressure is in reflect solar radiation, thus increasing the albedo;
agreement with geological findings (von Paris in contrast, high clouds, in particular ice-crystal
et al. 2008). cirrus clouds, add to atmospheric greenhouse
Faint Young Sun Paradox 841
warming, potentially sufficient to solve the FYSP luminosity integrated over the electromagnetic
(Rondanelli and Lindzen 2010) although full cov- spectrum) of main-sequence stars scales approx-
erage by an unrealistically thick and cold cloud imately with stellar mass to the third power;
cover would be required (Goldblatt and Zahnle further, the radius of the Earth’s orbit would
2011a). have been smaller for a more massive Sun (the
There is some evidence that elevated cosmic- orbital radius scales inversely with the solar
ray fluxes have a cooling effect on the Earth’s mass). Both effects combine to a scaling of the
atmosphere because they ionize tropospheric received flux with the fifth power of the
layers, and charged ion clusters lead to conden- solar mass.
sation nuclei that form reflective clouds (Shaviv Constraints on this model come from the
2003). But because the ionized wind of the young requirement that the young Martian atmosphere
Sun is thought to have been much stronger than was warm enough to maintain water in liquid F
today, the cosmic-ray flux reaching the inner form, but the Earth’s atmospheric temperature
solar system was more strongly suppressed, was moderate enough to prevent loss of the
therefore suppressing cloud formation in the water oceans in a runaway greenhouse. The
young Earth’s atmosphere and in turn leading to appropriate mass range for the young Sun was
a warmer climate (Shaviv 2003). This model has thus 1.03 MJ to 1.07 MJ. Corresponding solar
been observationally refuted, however (e.g., models are in acceptable agreement with
Bailer-Jones 2009 and references therein). helioseismology results (Sackmann and
A lower surface albedo may also be the result Boothroyd 2003).
of the suggested smaller continental area in early The observational evidence is presently
epochs although feedback on cloud coverage may unclear. Upper limits to the ionized-wind mass-
result in opposite trends. The absence of cloud loss rates deduced from radio observations of
condensation nuclei induced biologically young solar analogs are marginally compatible
(Charlson et al. 1987) also decreases the albedo. with the “bright young Sun” requirement, indi-
Considering these effects and claiming less cating a maximum zero-age main-sequence
reflective clouds because of larger cloud droplets (ZAMS) solar mass of 1.06 MJ (Gaidos
(because of fewer biogenically induced conden- et al. 2000). Indirect observations suggest a
sation nuclei), Rosing et al. (2010) inferred a smaller mass loss. If the solar wind had been
clement climate for the young Earth without the constant throughout the Sun’s main-sequence
need for high amounts of greenhouse gases life, then, including the losses from conversion
although this solution was again questioned of matter to energy in the hydrogen-burning core,
(Goldblatt and Zahnle 2011b). a total mass loss of only 0.0004 MJ would have
The amount of ice coverage and thus effects been achieved, which alters the solar radiative
on the energy balance can be influenced by var- output insignificantly. However, observations of
iations in the Earth’s axial tilt or its rotation “hydrogen walls” in the astrospheres of neigh-
period (slowly increasing with time due to the boring solar-like stars support an increased wind-
Moon’s tidal forces). Models for these effects mass loss in younger stars (Wood et al. 2005).
are inconclusive (Feulner 2012 and references A power-law trend extrapolated back to the
therein). ZAMS indicates a total mass loss of order
0.01 MJ, but some evidence shows that the
A Brighter Young Sun mass-loss rate was suppressed during the youn-
A radical remedy of the FYSP would be a young gest epochs; including this latter effect leads to a
Sun that was significantly more massive than at total mass loss of only 0.003 MJ (Minton and
present, losing the excess mass during its main- Malhotra 2007), again too little for a significant
sequence life in an enhanced ionized wind luminosity effect. The Wood et al. method has its
(Whitmire et al. 1995; Sackmann and Boothroyd own limitations, however, and strongly depends
2003). The bolometric luminosity (i.e., the on numerical modeling of the complex
interaction between magnetic stellar winds and Kasting JF (1991) CO2 condensation and the climate of
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Acad Sci U S A 106:14784–14789 netic emissions into space or by engineering its
Valley JW, Peck WH, King EM, Wilde SA (2002) A cool atmosphere to suppress the buildup of
early Earth. Geology 30:351–354 biosignature gases). Second is the case where
Von Paris P, Rauer H, Greenfell JL, Patzer B, Hedelt P,
Stracke B, Trautmann T, Schreier F (2008) Warming physical or/and chemical processes interfere,
the early earth – CO2 reconsidered. Planet Space Sci degrade, or hide (spectral) biosignature(s), e.g.,
45:1254–1259 via the presence of an optically thick cloud layer
Walker JCG, Hays PB, Kasting JF (1981) A negative which traps radiation below the cloud base or via
feedback mechanism for the long-term stabilization
of the Earth’s surface temperature. J Geophys Res photochemical processes which remove
86:9776–9782 biosignature gases in situ in the atmosphere, F
Whitmire DP, Doyle LR, Reynolds RT, Matese JJ e.g., via photolytic destruction in high UV envi-
(1995) A slightly more massive young Sun as an ronments. Third is the case where our (Earth-
explanation for warm temperatures on early Mars.
J Geophys Res 100:5457–5464 based) technology is insufficiently sensitive. For
Wood BE, M€uller H-R, Zank GP, Linsky JL, Redfield example, the microbial biosignature, nitrous
S (2005) New mass-loss measurements from Astrospheric oxide (N2O) while strongly attributable to life,
Lya absorption. Astrophys J Lett 628:L143–L146 has rather weak spectral absorption features,
which could be overlooked (related to this point,
see also the entry on the retrieval of atmospheric
properties of exoplanets).
Faint Young Sun Problem
▶ Faint Young Sun Paradox

See Also
▶ Bioindicator
▶ Exoplanetary Atmospheric Retrieval
False Negative
▶ False Positive
John Lee Grenfell
German Aerospace Center (DLR), Berlin,
Germany
False Positive
Definition John Lee Grenfell

German Aerospace Center (DLR), Berlin,
In general, a “false negative” refers to the result Germany
of an experiment or test which fails to support a
particular hypothesis, even though the hypothesis
is in fact correct. In an astrobiological context, Definition
“false negative” refers to the absence of a detect-
able life signal even though life is in fact present. A “false positive” refers to evidence which seem-
ingly supports a particular hypothesis or test but
which in fact results from unrelated causes. In an
Overview astrobiological context, “false positive” refers to
evidence which suggests the existence of life but
It is convenient to consider three cases when which in reality arises due to abiotic processes
searching for evidence of life. First is the case which are merely mimicking life.
844 Far Infrared (Far IR)
Overview
Fatty Acids, Geological Record of
Scientific validation of a proposed biosignature
requires the assessment and elimination of all Jennifer Eigenbrode
known abiotic (nonlife) processes (known as NASA Goddard Space Flight Center, Greenbelt,
the “false positives”) which could also produce MD, USA
such a signal. There are essentially two
issues: first to elucidate whether the species asso-
ciated with the signal (e.g., in the case of atmo- Keywords
spheric constituents such as ozone, methane, and
oxygen) is being produced by life or by Carboxylic acids; Cellular membranes; Lipids;
abiotic processes (this point is discussed further Molecular fossils; Organic acids
in the ▶ Bioindicator entry), and second, to
investigate whether the signal itself (e.g., the
atmospheric spectral band) is uniquely attribut- Synonyms
able to a biosignature gas, i.e., one can rule out or
subtract the effects of (partially) overlapping Aliphatic carboxylic acids; Alkanoic acids
spectral bands arising from other gas-phase
species, as well as interference from other phe-
nomena, e.g., the presence of clouds, pressure Definition
broadening, etc.
Fatty acids are ▶ carboxylic acids with an ali-
phatic tail (chain) that may be branched, satu-
rated, or unsaturated (i.e., contain double
See Also
bonds). Fatty acids typically have C12–C36 chain
lengths, though chains of only four carbons are
▶ Bioindicator
considered “fatty.” Fatty acids are constituents in
▶ False Negative
a variety of biomolecules and are released by the
hydrolysis of the ester linkages. Fatty acids are
important components of cellular ▶ membranes
of ▶ bacteria and eukaryotes, regulating both flu-
Far Infrared (Far IR) idity and permeability, and are key energy stores.
Fatty acids have also been observed in carbona-
▶ Infrared Astronomy ceous meteorites.
Overview
Farbstreifen Sandwatt In sediments and rocks, fatty acids may exist as

free fatty acids (i.e., hydrolyzed form) or
▶ Microbial Mats bound into larger biomolecules or geomolecules
(as in macromolecular ▶ kerogen) or bound to
minerals. Fatty acids have varying specificity
for particular phylogenetic groups of bacteria
and eukaryotes and also environmental
Fatty Acids conditions.
Most natural fatty acids have an even number
▶ Carboxylic Acids, Geological Record of of carbon atoms because they are biosynthesized
Fayalite 845
by the multiple additions of C2-acetyl units via ▶ Carbonaceous Chondrites, Organic Chemistry
the acetyl-CoA coenzyme. Organisms also of
biosynthesize a limited range of chain lengths ▶ Carbonyl
and have varying degrees of unsaturation ▶ Carboxylic Acid
(particularly plants, algae, and bacteria). ▶ Carboxylic Acids, Geological Record of
Biologically produced stereoisomeric configura- ▶ Cell Wall
tions of unsaturated fatty acids are all cis ▶ Complex Organic Molecules
(the Latin term cis describes the orientation of ▶ Dicarboxylic Acid
functional groups, on the same side, within a ▶ Eukarya
molecule; the opposite term is trans, meaning ▶ Hydroxy Acid
“on the other side” or “across”). All of these ▶ Kerogen
features are regarded as general biosignatures ▶ Membrane F
and are not observed in abiological fatty acids, ▶ Molecular Fossils
such as those observed in meteorites or in
synthetics.
Fatty acid abundance diminishes during early
diagenesis primarily due to in situ microbial References and Further Reading
reworking. Preferential loss of unsaturated and
Eigenbrode JL (2007) Fossil lipids for life-detection: a
short-chain moieties is commonly observed. case study from the early Earth record. Space Sci
However, in the presence of other refractory Rev 135:161–185
materials, fatty acids will be incorporated into Killops S, Killops V (2005) Introduction to
geomolecules that enhance preservation. Satu- organic geochemistry, 2nd edn. Blackwell, Oxford,
p 393
rated, o-hydroxyl, and a,o-diacids are more Meyers PA, Leenheer MJ, Bourbonniere RA (1995) Dia-
resistant to diagenesis and have been observed genesis of vascular plant organic matter components
in the ancient geological record. A variety of during burial in lake sediments. Aquat Geochem
reactions can lead to the loss of carbon from the 1:35–52
Petsch ST, Eglinton TI, Edwards KJ (2001) 14C-dead
alkyl chain, which during later diagenesis leads living biomass: evidence for microbial assimilation
to molecular patterns having greater relative of ancient organic carbon during shale weathering.
abundance of odd-carbon chain lengths. For Science 292:1127–1131
example, decarboxylation of even-carbon chain Petsch ST, Edwards KJ, Eglinton TI (2003) Abundance,
distribution and d13C analysis of microbial
length fatty acids leads to odd-carbon alkane phospholipid-derived fatty acids in a black shale
chains. Further, thermal maturation and carbon weathering profile. Org Geochem 34:731–743
loss of alkanes lead to a diminishing preference Petsch ST, Edwards KJ, Eglinton TI (2005)
for odd or even chain lengths. Thus, in the rock Microbial degradation of sedimentary organic
matter. Palaeogeogr Palaeoclimatol Palaeoecol
record, molecular distributions of fatty acids and 219:157–170
alkanes can indicate biological sources and a Sephton MA (2005) Organic matter in carbonaceous
relative degree of diagenesis. In some cases, meteorites: past, present and future research. Philos
fatty acids from microbes living in rocks or petro- Trans R Soc A 363:2729–2742
Summons RE, Albrecht P, McDonald G, Moldowan JM
leum reservoirs can complicate molecular inter- (2007) Molecular biosignatures: generic qualities of
pretations and are important records of recent organic compounds that betray biological origins.
processes. Space Sci Rev 135:133–157
See Also
▶ Acid Hydrolysis Fayalite

▶ Bacteria
▶ Biomarkers ▶ Olivine
846 Feeding Zone
Feeding Zone Fennoscandia
Sean N. Raymond Pentti Hölttä

Laboratoire d’Astrophysique de Bordeaux, Department of Geosciences and Geography,
CNRS, Universite de Bordeaux, France University of Helsinki, Finland and Geological
Survey of Finland, Espoo, Finland
Definition Keywords
A planet’s feeding zone is the region within the Archean; Fennoscandia; Greenstone belt; TTG
▶ protoplanetary disk from where solids can be
gravitationally captured by the planet. The plan-
etary feeding zone is schematically a ring in
the protoplanetary disk, centered on the Synonyms
central star, the forming planet being located in
the middle of the ring. The size of the feeding Fennoscandian shield
zone depends on different factors, in particular
the planetary mass, its distance to the central star,
and the dynamical state of the solids. Typically,
the size of the feeding zone is of the order of a few Definition
times the Hill Radius of the planet.
The size and location of the feeding zone has Fennoscandia is the geographic term that covers
an influence on both the formation process of the Scandinavian countries, Finland, and the
the planet (and therefore the final mass of a northwesternmost part of Russia. The geologic
planet) and its composition. Large feeding term Fennoscandian Shield denotes the Precam-
zones, which contain a lot of mass, are brian units of this area. The Precambrian of
required for giant planet formation. Given that Fennoscandia consists of Archean and Protero-
radial temperature gradients are thought to trans- zoic rocks, this description dealing with the
late into compositional variations in the proto- former.
planetary disk, a planet’s feeding zone has
consequences for its composition. A narrow feed-
ing zone implies that the planet’s composition
should represent material condensed within a Overview
small range in temperatures, and a wide feeding
zone implies significant radial mixing during Archean rocks comprise much of the eastern and
accretion. northern parts of the Fennoscandian shield. They
have been divided into three major provinces: the
Karelian Province in the west, the Belomorian
Province in the east, and the Kola province in
See Also the north. Neoarchean and Mesoarchean
(3.2–2.7 Ga) lithologies prevail, but a high pro-
▶ Planetesimals portion fall in a relatively restricted range of
▶ Protoplanetary Disk ca. 2.84–2.67 Ga. Older gneisses of Paleoarchean
Fennoscandian Shield 847
age (ca. 3.5 Ga) have been documented in a small See Also
area in the western part of the Karelian Province,
and Mesoarchean 3.2–2.9 Ga rocks occur in ▶ Amphibolite Facies
restricted areas in the western and eastern parts ▶ Archean Eon
of the Karelian Province. A ▶ tonalite- ▶ Banded Iron Formation
trondhjemite-granodiorite (TTG) association ▶ Craton
with subordinate ▶ greenstone belts, ▶ Earth, Formation and Early Evolution
paragneisses, granulite complexes, and ▶ Greenstone Belts
migmatitic amphibolites dominates the Archean ▶ Igneous Rock
terranes. The TTG magmatism mainly occurred ▶ Metamorphic Rock
between 2.83 and 2.74 Ga and was followed by a ▶ Metasediments
brief period of sanukitoid magmatism between ▶ Ophiolite F
2.73 and 2.70 Ga. Many TTG have adakitic geo- ▶ Shield
chemical signatures. Sanukitoids originate from ▶ Tonalite-Trondhjemite-Granodiorite
melting of a metasomatized mantle source, prob-
ably as a result of a slab break-off following a
continental collision (Halla et al. 2009). The
youngest 2.71–2.69 Ga granites are thought to References and Further Reading
result from melting of the crust during collisional
processes. The youngest Archean igneous rocks Halla J, van Hunen J, Heilimo E, Hölttä P (2009) Geo-
chemical and numerical constraints on Neoarchean
are 2.67–2.61 Ga anorogenic alkaline granitoids plate tectonics. Precambrian Res 174:155–162
that occur in the Keivy terrane in Kola (Zozulya Shchipansky AA, Samsonov AV, Bibikova EV, Babarina
et al. 2005), the 2.61 Ga Siilinjärvi carbonatite II, Konilov AN, Krylov KA, Slabunov AI, Bogina MM
close to the western border of the Karelian (2004) 2.8 Ga boninite-hosting partial
suprasubduction ophiolite sequences from the North
Province, and some 2.61 Ga mafic dykes in the Karelian greenstone belt, NE Baltic Shield, Russia. In:
eastern part of the Karelian Province (Slabunov Kusky T (ed) Precambrian ophiolites and related
et al. 2006). rocks. Elsevier, Amsterdam, pp 425–487
The oldest (ca. 3.10–2.90 Ga old) volcanic Slabunov AI, Lobach-Zhuchenko SB, Bibikova EV,
Sorjonen-Ward P, Balagansky VV, Volodichev OI,
rocks are found in greenstone belts in the eastern Shchipansky AA, Svetov SA, Chekulaev VP, Arestova
and northwestern parts of the Karelian Province. NA, Stepanov VS (2006) The archaean nucleus of the
Most greenstone complexes are 2.88–2.78 Ga Baltic/Fennoscandian shield. In: Gee DG, Stephenson
old, often showing an assembly of plume-related RA (eds) European lithosphere dynamics. Geol Soc
London, Memoir 32. pp 627–644
komatiites and tholeiites, island arc-type calc- Volodichev OI, Slabunov AI, Bibikova EV, Konilov AN,
alkaline volcanic rocks, as well as Kuzenko T (2004) Archaean eclogites in the
▶ metasediments and ▶ banded iron formations. Belomorian mobile belt, Baltic shield. Petrology
Neoarchean 2.72 Ga-old eclogites metamor- 2:540–560
Zozulya DR, Bayanova TB, Eby GN (2005) Geology and
phosed at 14–17 kbars are known in a few areas age of the late Archean Keivy alkaline province,
in the Belomorian Province (Volodichev Northeastern Baltic Shield. J Geol 113:601–608
et al. 2004). The Seriak and Iringora greenstone
belts in the Belomorian Province have
▶ ophiolite-like features (Shchipansky
et al. 2004). Greenstone belts in the central parts
of the Karelian Province are younger Fennoscandian Shield
(2.75–2.73 Ga) than in its western and eastern
parts. ▶ Fennoscandia
848 Ferment (Obsolete)
Overview
Ferment (Obsolete)
Many microorganisms, obligate or facultative
▶ Enzyme anaerobic, are able to degrade extracellular poly-
mers (e.g., polysaccharides, proteins, nucleic
acids) and use the monomers (e.g., hexoses, pen-
toses, amino acids, purines, pyrimidines) as fer-
Fermentation mentable substrates. There are other substrates of
fermentations such as organic acids (e.g., citrate,
Juli Peretó succinate, or malonate) or even aromatic hydro-
Institut Cavanilles de Biodiversitat i Biologia carbons. In this latter case, fermentation appears
Evolutiva, Universitat de València, València, as a metabolic task of several microbial species
Spain working together. Regarding amino acid fermen-
tations, many anaerobic bacteria can catabolize
single amino acids but grow better with amino
Definition acid mixtures (the Stickland reaction). In this
case, some amino acids of the mixture act as
Fermentation is an anaerobic ▶ catabolism of a reductants, whereas others are the oxidants.
reduced carbon source (e.g., glucose) to generate The stoichiometric yield of ATP in a fermen-
▶ ATP within a strict internal ▶ oxidation- tation depends on the particular pathway used and
▶ reduction balance. In many cases, the same can range from less than one up to 4 mol of ATP
substrate is used both as reductant and oxidant. per mol of fermentable substrate. Microorgan-
The hallmark of fermentation is the accumulation isms are capable of generating a wide array of
of partially oxidized end products. Only a part of end products during fermentation, including car-
the chemical energy stored in the initial substrate bon dioxide, ethanol, lactate, butyrate, acetate,
is conserved during the synthesis of ATP, usually and propionate. In comparison to respiration
by a mechanism of substrate-level phosphoryla- (i.e., electron transport chain-dependent pro-
tion. Nevertheless, there are some examples of the cesses), fermentations are less energetically effi-
involvement of an electrochemical ion gradient in cient because a lot of potential chemical energy is
the synthesis of ATP (e.g., citrate fermentation). still retained in most of the end products. Thus, to
compensate this relatively low-energy yield,
large amounts of fermentable substrate are used
History and most carbons from this can be recovered in
the form of end products. Other anaerobic bacte-
Antoine Laurent Lavoisier (1743–1794) col- ria use the excreted end products of fermenta-
lected the first quantitative data on alcoholic fer- tions, building an anaerobic food chain with
mentation. Louis Pasteur (1822–1895) studied in methanogens at the bottom. From a biotechno-
depth the fermentation by intact living cells and logical point of view, however, since very ancient
defined this physiological process as “life without times, fermentations are widely used for the
oxygen.” In 1897, Eduard Buchner (1860–1917) processing of food products, such as yogurt,
showed that fermentation could occur in cell-free cheese, beer, wine, or bread.
extracts, inaugurating the genuine biochemical In a few cases, some ATP synthesis during
in vitro approach to living phenomena. Parallel fermentation is associated with the dissipation
studies on yeast alcoholic fermentation and of an electrochemical potential gradient either
anaerobic muscle glycolysis during the first half of protons or sodium ions. These ion gradients
of the twentieth century contributed to the devel- can be generated by electron transport (e.g.,
opment of biochemistry as a science (Barnett fumarate reduction in Propionibacterium), mem-
2003). brane decarboxylases (e.g., sodium-pumping
Fermi Paradox 849
succinate decarboxylase of Propionigenium Definition

modestum), or electrogenic substrate transloca-
tion through membranes (e.g., lactic acid bacteria The Fermi paradox, attributed to Italian
like Lactococcus cremoris). physicist Enrico Fermi, concerns the apparent
When compared to the other energy- contradiction between the lack of any
generating systems (such as respiration and pho- evidence for the presence of extraterrestrials
tosynthesis), fermentation seems simpler at the on Earth and the view that extraterrestrial
structural and enzymatic levels. For this reason, civilizations should be rather common in the
in 1924, Oparin (1924) postulated that fermenta- Galaxy.
tion was the earliest metabolic mode.
History F
See Also
Fermi formulated his paradox in 1950, during a
▶ Anaerobe casual conversation in Los Alamos Laboratory
▶ ATP with E. Teller and colleagues, with the famous
▶ Catabolism phrase “where are they?” (or “where is every-
▶ Chemoorganotroph body?”). His point was that, if there are many
▶ Embden-Meyerhof-Parnas Pathway extraterrestrial civilizations, Earth should have
▶ Glycolysis been visited by one or more of them, long ago
▶ Origin of Life and many times over. The discussion went
▶ Oxidation completely unnoticed for many years. The phrase
▶ Reduction “where are they?” attributed to Fermi but without
comments is first encountered in a paper
published in 1963 by American astronomer Carl
Sagan. Sagan referred to this problem as “Fermi’s
Barnett JA (2003) A history of research on yeasts 5: the paradox” after American astronomer Michael
fermentation pathway. Yeast 20:509–543 Hart independently rediscovered Fermi’s argu-
Kim BH, Gadd GM (2008) Bacterial physiology and ments in 1975.
metabolism. Cambridge University Press, Cambridge,
Chapter 8
Oparin AI (1924) Proiskhozhedenie Zhizni. Mosckovskii
Rabochii, Moscow (Reprinted and translated in Bernal Overview
JD (1967) The origin of life. Weidenfeld and Nicolson,
London) As all paradoxes, the Fermi paradox is better
White D (1995) The physiology and biochemistry of pro-
karyotes. Oxford University Press, New York, Chapter 13 understood if its premises are explicitly stated:
1. There are many extraterrestrial civilizations in

the Milky Way.
2. Our civilization is a typical one (not the first to
Fermi Paradox have appeared, neither the most technologi-
cally advanced, nor the only one seeking to
Nikos Prantzos explore the cosmos and communicate with
Institut d’Astrophysique de Paris, Paris, France others).
3. Interstellar travel is not too difficult
for civilizations slightly more advanced
Keywords than ours.
4. Galactic colonization (either by some of those
Extraterrestrial civilizations civilizations or their self-replicating robots)
850 Fermi, Enrico
can be achieved in less than 108 years, i.e., less References and Further Reading
than 1 % of the age of the Galaxy.
Prantzos N (2000) Our cosmic future – humanity’s fate in
the universe. Cambridge University Press, Cambridge
If hypotheses (a) to (d) are valid, the conclu-
Prantzos N (2013) A joint analysis of the Drake equation
sion “they should be here” is logically deduced, and the Fermi paradox. Int J Astrobiol 12:246–253
and the Fermi paradox applies. It ceases to apply Webb S (2002) Where is everybody? Fifty solutions to the
if one or more of its premises are refuted. Fermi paradox. Copernicus Books – Praxis, Chichester
Supporters of ETI (extraterrestrial intelligence)
reject one (or more) of the assumptions (b) to (d),
in order to save the key hypothesis (a). In con-
trast, opponents of ETI uphold the plausibility of Fermi, Enrico
(c) and (d), while rejecting (b) and even (a).
For instance, British astronomer Fred Hoyle Fernando B. Figueiredo
thought that interstellar distances make interstel- CITEUC, University of Coimbra, Coimbra,
lar travel impossible. However, arguments most Portugal
often discussed refer to the sociological
rather than the physical aspects of the problem.
Fermi believed that a technological civilization History
would be too short-lived, destroying itself
before mastering interstellar travel. Long Enrico Fermi (1901–1954) was an Italian physi-
before him, the Russian savant and father cist famous by his important contributions to the
of astronautics Konstantin Tsiolkovsky knowledge and development of the quantum
preferred the so-called “zoo hypothesis” world, mainly on the nuclear and particle physics.
(reformulated independently in 1973 by Ameri- In 1926, Fermi discovered, independently of Paul
can astronomer John Ball), according to which Dirac, the statistical laws, nowadays known as
advanced civilizations observe us from afar with- the Fermi-Dirac statistics, governing the particle
out interfering, for various reasons (e.g., waiting systems subject to Pauli’s exclusion principle
for us to gain “maturity” before joining the (the fermions).
“cosmic club”). Between 1934 and 1936, he published two
All sociological arguments share a common important papers on the artificial radioactivity
weak point: It is hard to believe that any one of produced by neutron bombardment and on the
them applies to every single civilization in the absorption and diffusion of slow neutrons (this
Galaxy. Moreover, in such a case, assumption one led to the discovery of nuclear fission). In
(b) would be implicitly violated since we would 1938, he won the Nobel Prize in Physics for his
be the only civilization seeking communication work on those issues.
with others. The most “economic” solution to In 1939, Fermi and his family, his Jewish wife
Fermi’s paradox consists in straightforwardly Laura Capon and their two children, Nella and
rejecting hypothesis (a): Technologically Giulio, emigrated to the USA, where he worked
advanced civilizations are rare in the Galaxy in the Manhattan Project during World War II. In
(and, perhaps, we are alone). 1950, while working at the Los Alamos Labora-
tory, Fermi and some colleagues discussed about
the possibility of existence of non-terrestrial
intelligent life. Some of them supported the idea
See Also that due to the apparent size and age of the Uni-
verse, the probability of having been developed
▶ Drake Equation alien civilizations would be huge. And if so the
▶ Fermi, Enrico aliens have had more than enough time to travel
▶ Galactic Habitable Zone around the Galaxy and visit the Earth. This
Fig Tree Group 851
prompted Fermi to ask, where are they? This in viral RNA polymerases and reverse transcrip-
significant issue became known as the Fermi par- tases results in low-fidelity replication, with aver-
adox. In the 1980s, there is a massive scientific age error rates of 103–105 mutations per
production of writings by scientists in and out of nucleotide and round of copy.
the SETI community to address the Fermi
paradox.
The NASA’s Fermi Gamma-ray Space Tele-
See Also
scope was launched in 2008 with the scientific
mission to study high-energy phenomena and the
▶ Error rate
exotic gamma-ray sources, from ancient black
▶ Mutation
holes, to rapidly rotating neutron stars, or pulsars,
▶ Replication (Genetics)
in the Milky Way galaxy, to jets powered by F
▶ Template
supermassive black holes in far-away young
galaxies.
Fig Tree Group

See Also
Christoph Heubeck
▶ Fermi Paradox
Institut f€ur Geowissenschaften, Friedrich-
▶ SETI
Schiller-Universität Jena, Jena, Germany
Keywords
Fidelity
Archean ocean; Barite; Chert; Banded-iron for-
Carlos Briones mation; Early life
Spain Definition
The Fig Tree Group overlies the Onverwacht

Synonyms Group and is overlain by the Moodies Group in
the Paleoarchean ▶ Barberton Greenstone Belt,
Replication accuracy South Africa and Swaziland. Strata of this
group were deposited between approximately
3,258 3 and 3,225 3 Ma, reach up to 3 km
Definition in thickness, and show a wide variety of deposi-
tional settings and lithologies.
In molecular biology and genetics, fidelity is the
accuracy of template-dependent nucleic acid
copying, in particular DNA or RNA replication. Overview
Since accurate genome replication is critical for
the viability of cellular organisms, proofreading The Fig Tree Group consists largely of
mechanisms ensure near-perfect fidelity for DNA interbedded siliciclastic and chemically precipi-
replication, with average error rates of tated sedimentary strata, deposited in a range of
109–1012 mutations per nucleotide and round settings, as well as intermediate to felsic volca-
of copy. In turn, the lack of proofreading activity nics. In contrast, underlying strata of the
852 Fig Tree Group
▶ Onverwacht Group are dominated by mafic tectonic setting. Subsequent deformation, how-
and ultramafic volcanics, with very little terrige- ever, has overprinted many of the original
nous contribution; strata of the overlying depositional relationships. Lithology, prove-
▶ Moodies Group consist largely of shallow- nance, depositional environment, volcanism,
water quartz-rich sandstones. Hofmann (2005) and style of deformation of the Fig Tree
and Lowe and Byerly (2007) provide reviews of Group are compatible with a tectonic setting
the stratigraphy of the Fig Tree Group. analogous to a modern convergent margin, such
Aside from thick fan-delta conglomerates, as a retroarc foreland, interarc, or forearc basin.
turbiditic greywackes, and fine-grained ferrugi- In that regard, the Fig Tree strata record a transi-
nous and tuffaceous strata (Heinrichs 1980), the tion from the oceanic-plateau setting of the
Fig Tree Group south of the Inyoka Fault includes underlying Onverwacht Group to shallow-water
thick jaspilitic ▶ banded-iron formation (within continental settings of the overlying Moodies
the Ngwenya Formation of Hofmann 2005), mul- Group.
tiple sedimentary barite beds (in the lower
Mapepe Formation near the base of the Group),
banded carbonaceous ▶ cherts, and several – in See Also
part reworked – impact-derived spherule beds.
Fig Tree strata north of the Inyoka Fault largely ▶ Archean Environmental Conditions
consist of deeper-water turbiditic siliciclastic ▶ Archean Tectonics
rocks. Northern and southern facies are overlain ▶ Banded Iron Formation
by dacitic volcanic and volcaniclastic rocks of the ▶ Barberton Greenstone Belt
Schoongezicht Formation (Condie et al. 1970) ▶ Barberton Supergroup
and the Auber Villiers Formation, respectively ▶ Barite
(Lowe and Byerly 1999). ▶ Moodies Group
Together, these lithologies provide a wealth of ▶ Onverwacht Group
unique insights in the chemical and physical set- ▶ Spherules
ting of the Archean ocean, demonstrating the
interplay of organic matter, dissolved Fe, Si,
and Ba, and the temporary presence of
(photolytically generated?) sulfate (Knauth and Bao H, Rumble D, Lowe DR (2007) The five stable iso-
Lowe 2003; Bao et al. 2007; Roerdink tope compositions of Fig Tree barites: implications on
et al. 2013). The volcanic strata and hypabyssal sulfur cycle in ca. 3.2 Ga oceans. Geochim
intrusions in the Fig Tree Group are contempora- Cosmochim Acta 71:4868–4879
Condie KC, Macke JR, Reimer TO (1970) Petrology and
neous with TTG plutons adjacent to the BGB, geochemistry of early Precambrian greywackes from
adding another tectonic perspective to our under- Fig Tree Group, South Africa. Geol Soc S Afr Bull
standing of this unit. 81:2759–2776
The Fig Tree Group developed in a highly Heinrichs T (1980) Lithostratigraphische Untersuchungen
in der Fig Tree Gruppe des Barberton Greenstone Belt
structured paleotopography, including terrestrial zwischen Umsoli und Lomati (S€ udafrika). Göttinger
alluvial fan deltas, above-wavebase platforms, Arb Geol Paläontol 22:118
shelf bank settings, and various deepwater facies. Hofmann A (2005) The geochemistry of sedimentary
The reasons for the complex paleotopography rocks from the Fig Tree Group, Barberton greenstone
belt: implications for tectonic, hydrothermal and sur-
may relate to the setting at the margin of the face processes during mid-Archaean times. Precam-
Onverwacht volcanic plateau, syndepositional brian Res 143:23–49
deformation related to major thrusting (early D2 Knauth LP, Lowe DR (2003) High Archean climatic tem-
of Lowe and Byerly, 2007), and the proximity to perature inferred from oxygen isotope geochemistry of
cherts in the 3.5 Ga Swaziland Supergroup, South
the Inyoka Fault system. Lowe (2013) suggested Africa. Geol Soc Am Bull 115:566–580
that major meteorite impacts (recorded by spher- Lowe DR (2013) Crustal fracturing and chert dike forma-
ule beds) may have contributed to the change in tion triggered by large meteorite impacts, ca. 3.260 Ga,
Fischer-Tropsch Effects on Isotopic Fractionation 853
Barberton greenstone belt, South Africa. Geol Soc Am Vertical lines lie either in or behind the plane of
Bull 125:894–912 the paper, while horizontal lines project out from
Lowe DR, Byerly GR (1999) Stratigraphy of
the west-central part of the Barberton the surface. In a Fischer projection, the plane of
Greenstone Belt, South Africa. Geol Soc Am Spec symmetry (3D) between two enantiomers
Pap 329:1–36 becomes a line of symmetry between their 2D
Lowe DR, Byerly GB (2007) An overview of the representations.
geology of the Barberton Green Belt and vicinity:
implications for early crustal development. In:
Van Kranendonk MJ, Smithies RH, VC Bennett
(eds) Earth’s oldest rocks. Developments in
precambrian geology, Elsevier, Amsterdam, 15. See Also
pp 481–526
Roerdink DL, Mason PRD, Whitehouse MJ, Reimer
T (2013) High-resolution quadruple sulfur isotope ana- ▶ Carbohydrate F
lyses of 3.2 Ga pyrite from the Barberton Greenstone ▶ Chirality
Belt in South Africa reveal distinct environmental
controls on sulfide isotopic arrays. Geochim
Cosmochim Acta 117:203–215
Fischer-Tropsch Effects on Isotopic

Fischer Projection Fractionation
Henderson James (Jim) Cleaves II Daniele L. Pinti

Earth–Life Science Institute (ELSI), Tokyo GEOTOP Research Center for Geochemistry and
Institute of Technology, Meguro–ku, Tokyo, Geodynamics, Université du Québec à Montréal,
Japan Montréal, QC, Canada
USA
Blue Marble Space Institute of Science, Keywords
Washington, DC, USA
Center for Chemical Evolution, Georgia Institute Carbon isotopes; Hydrothermal environments;
of Technology, Atlanta, GA, USA Isotope biological markers; Metal-catalyzed
reactions
Definition
Definition
In chemistry, a Fischer projection is a
two-dimensional representation of a three- The Fischer-Tropsch-type reaction is a method
dimensional organic molecule devised by Emil for the synthesis of hydrocarbons and some ali-
Fischer in 1891. Fischer projections depict the phatic compounds in the presence of a metal
stereochemistry of molecules and are thus useful catalyst. The carbon isotopic ratios of produced
for depicting enantiomers of chiral molecules, hydrocarbons (methane and short-chain hydro-
especially monosaccharides. The carbon back- carbons) and byproducts such as carbon dioxide
bone of the molecule is depicted as a vertical show large fractionation relative to the reactants.
line and the bonds of side chains are represented Abiotic organic products are depleted in 13C to a
as horizontal lines, with carbon atoms degree typically ascribed to biological processes,
represented by the center of crossing lines. The indicating that carbon isotopic composition may
orientation of the carbon chain is such that the C1 not be a particularly effective discriminant
carbon is at the top of the molecule diagrammed. between biologic and nonbiologic sources.
854 Fischer-Tropsch Effects on Isotopic Fractionation
Overview 50‰, overlapping typical d13C values of

thermogenic CH4. McCollom and Seewald
▶ Fischer-Tropsch-type reactions (FTT hereaf- (2006) produced H2, CO2, CH4, and light hydro-
ter) can produce various light and heavy hydrocarbons (C2–C5) by heating an aqueous solution
carbons, oxygen-containing compounds of ▶ formic acid (HCOOH) in the presence of
(▶ alcohols, ▶ carboxylic acids), and basic iron powder at 250 C and 325 bar in a hydro-
N-compounds, including ▶ amino acids thermal cell. The d13C of the dissolved CO2 was
(Hayatsu and Anders 1981). It is now widely 14‰ and values for methane and light hydro-
accepted that at mantle and crustal temperatures carbons were as low as 49‰. Again these
and pressures and in presence of natural metal results show that organic products are depleted
catalysts such as iron, FTT can produce carbona- in 13C to a degree typically ascribed to biological
ceous matter. Of particular interest for astrobiol- processes, indicating that the carbon isotopic
ogists are FTT reactions in the seafloor composition may not be an effective diagnostic
▶ hydrothermal environments, this being one of means to differentiate between biologic and
the possible environments where Hadean or nonbiologic sources (McCollom and Seewald
Archean life could have originated and evolved. 2006).
The carbon isotopic compositions of the car-
bonaceous products of FTT reactions display var-
ious extents of isotopic fractionation relative to
the reactant. The majority of products are See Also
depleted in 13C, with d13C values (where
d13C = [(13C/12C)sample/(13C/12C)standard 1] ▶ Biomarkers
and standard is Pee Dee Belemnite) showing ▶ Biomarker, Isotopic
from 2 to 60‰ (‰ = permil) depletion compared ▶ Carbon Isotopes as a Geochemical Tracer
to the reactant. Several investigators (e.g., Lancet ▶ Fischer-Tropsch-Type Reaction
and Anders 1970; Kerridge et al. 1989) synthe- ▶ Hydrothermal Environments
sized hydrocarbons from CO and H2 at ▶ Serpentinization
100–500 C in the presence of metal catalysts
(Fe, Co, Ni, ▶ magnetite, etc.) and proposed
that such a process could explain the
origin of carbonaceous and graphitic matter in References and Further Reading
meteorites. Their results indicated that short-
chain hydrocarbons (C1–C4) were usually more Hayatsu R, Anders E (1981) Organic compounds
in meteorites and their origins. Top Curr Chem
depleted in 13C than the reactant CO. However, 99:1–37
the carbon isotopic signature of experimentally Horita J (2005) Some perspectives on
produced methane (CH4) can vary significantly, isotope biosignatures for early life. Chem Geol
from very depleted values of 60‰ to values 218:171–186
Horita J, Berndt ME (1999) Abiogenic methane formation
even heavier than the reactant CO (Lancet and and isotopic fractionation under hydrothermal condi-
Anders 1970). tions. Science 285:1055–1057
Horita and Berndt (1999) conducted a series of Kerridge JF, Mariner R, Flores J, Chang S (1989) Isotopic
experiments to investigate isotopic fractionation characteristics of simulated meteoritic organic matter:
1. Kerogen-like material. Orig Life Evol Biosph
during hydrothermal production of CH4 from 19:561–572
CO2 and H2 in the presence of Ni-Fe alloy. Lancet MS, Anders EA (1970) Carbon isotope fraction-
These experiments simulate the production of ation in the Fischer-Tropsch synthesis and in meteor-
abiotic methane and other organic compounds ites. Science 170:980–982
McCollom TM, Seewald JS (2006) Carbon isotope com-
during ▶ serpentinization. The d13C values of position of organic compounds produced by abiotic
abiotic CH4 produced from dissolved CO2 with synthesis under hydrothermal conditions. Earth Planet
a mantle-like 13C signature varied from 4 to Sci Lett 243:74–84
Fischer-Tropsch-Type Reaction 855
liquid fuels from coal by indirect catalytic hydro-

Fischer-Tropsch-Type Reaction genation of carbon oxides (Hindermann
et al. 1993). FTT chemistry was a natural exten-
Natasha M. Johnson1 and Joseph Andrew Nuth III2 sion of the pioneering work of Sabatier and
1
NASA Goddard Space Flight Center, Greenbelt, Senderens (1902), who synthesized ▶ methane
MD, USA by passing a mixture of ▶ hydrogen and
2
Solar System Exploration Division, NASA’s CO/CO2 over a reduced nickel catalyst. It is
Goddard Space Flight Center, Greenbelt, MD, important to remember that formation of methane
USA (CH4) alone is not the Fischer-Tropsch reaction
but simply methanation. FTT synthesis was
widely used in Germany during World War II to
Keywords manufacture organic fuels in the form of gaso- F
line, diesel oil, liquefiable gases, and paraffin
Catalyst; Hydrocarbons; Hydrogenation; Macro- wax. Early FTT experimentation used nickel
molecular organics; Methanation; Organics and then iron catalysts. However, iron suffered
from excessive carbon deposition leading to
blockage of the ▶ catalyst pores and deactivation
Definition (Pearce et al. 1989). As a result, modern FTT
plants typically use precious metals, cobalt, or
In Fischer-Tropsch-type (FTT) synthesis hydro- nickel dispersed on substrates (alumina, silica,
carbons are produced through hydrogenating car- etc.). In actuality, many of these “fresh” catalytic
bon monoxide via surface-mediated reactions, as surfaces are oxides, and it is these metal oxides
shown in reaction (1): that have industrial catalytic properties.
nCO þ ð2n þ 1ÞH2 ! Cn H2nþ2 þ nH2 O (1)

Overview
The FTT catalytic sequence, as other forms of
catalysis, is rather complex and involves several The origins of organics in the early Solar System
related reactions on the catalyst surface. A typical are complex and still somewhat poorly understood
FTT experiment involves circulating gases (Ehrenfreund and Charnley 2000). However,
(typically CO and H2) through and/or over a Fischer-Tropsch-type (FTT) catalytic reduction of
catalyst at a specific temperature and then mea- CO by hydrogen to produce methane and other
suring the resulting gases (such as water, meth- ▶ hydrocarbons has long been recognized as an
ane, and carbon dioxide) and residue on the important potential source of organic material in
grains as reaction time progresses. As the grains the ▶ Solar Nebula, since the “basic ingredients”
became coated with residue, organics are proof this reaction (H2, CO, and plausible catalysts)
duced faster and in greater quantities and CO is were ubiquitous in this environment (Kress and
quickly depleted. The reaction eventually reaches Tielens 2001). Hayatsu and Anders (1981) com-
a plateau. Once this point is reached, the gases are pared the results of a wide range of experimental
removed and the system refilled to be run again. studies of FTT reactions to detailed analyses of the
organic residue extracted from ▶ carbonaceous
chondrites. They reported that a significant fraction
History of the longer-chain hydrocarbons and more com-
plex aromatic compounds in meteorites could be
Fischer-Tropsch-type synthesis is a common explained by FTT chemistry. However, there are
industrial process developed by coal researchers some notable exceptions. In particular, ▶ amino
Franz Fischer (1877–1947) and Hans Tropsch acids are not produced efficiently enough or in the
(1889–1935) for obtaining gasoline and other proportions and compositional distributions
856 Fischer-Tropsch-Type Reaction
observed in meteorites via FTT reactions. This is in These coatings are composed of macromolecular
contrast to the ▶ Miller-Urey synthesis, which has organic phases (Johnson et al. 2004, 2007). These
been successful in generating many of the amino experiments also showed that as the grains became
acids associated with living systems and carbona- coated, Haber-Bosch-type reactions (catalytic
ceous chondrites. reduction of N2 by hydrogen to make ammonia)
In experimental studies of FTT reactions, pulver- took place, resulting in nitrogen-bearing organics
ized iron-nickel meteorites, powdered carbonaceous (Hill and Nuth 2003).
chondritic meteorites, and synthetic substrate- These types of FTT reactions are an effective
supported nickel-iron particles have been used as means to produce complex hydrocarbons.
the active catalyst (Hayatsu and Anders 1981; Organics generated by this technique could rep-
Llorca and Casanova 2000). In astrochemical exper- resent the carbonaceous material incorporated in
iments designed to measure the efficiency of FTT comets and meteorites.
reactions (Hong and Fegley 1998; Fegley 1999),
metallic iron (or iron oxide) was also used as the
catalyst. These materials are relevant in that they Key Research Findings
were surely present in the primitive nebula. How-
ever, the bulk form of iron meteorites is not condu- Dust grains falling into a protostellar system can
cive to efficient catalysis, nor is there evidence that provide surfaces that promote the reaction of H2,
iron-nickel particles that might have been found in N2, and CO into both volatile organics and a
the nebula would have been analogous to powdered macromolecular coating that continues to pro-
iron meteorites. In addition to the abovementioned mote the formation of organic materials.
materials, preliminary experiments (Ferrante Although the reaction is most efficient in the
et al. 2000) demonstrated that iron silicate grain innermost regions of the nebula, this does not
analogues – even those containing 10 % total pose significant problems as the reaction products
iron – are effective FTT catalysts. as well as the coated grains can migrate back out
Amorphous iron silicates provide reasonably to the far reaches of the nebula, thus seeding the
good surfaces to promote the reaction of H2, N2, entire nebula with the organic building blocks of
and CO to organic materials, resembling those life (Nuth et al. 2008).
found in meteorites (Hill and Nuth 2003; Gilmour
et al. 2002). Although not as efficient, amorphous
magnesium silicates, bronzite, and a range of other See Also
natural materials also promote the formation of
organic materials from H2 and CO. Fortunately, ▶ Fischer-Tropsch Effects on Isotopic
the amorphous metal-silicate grains or “smokes” Fractionation
are straightforward to produce in the laboratory ▶ Miller, Stanley
and serve as good early Solar Nebula catalyst ▶ Protoplanetary Disk, Chemistry
analogues. In addition to producing volatile ▶ Solar Nebula
organics via the reaction of H2 and CO on these ▶ Urey’s Conception of Origins of Life
surfaces, a major reaction product appears to be a
macromolecular organic coating on the grain sur-
faces. However, there is no significant decrease in
the reaction rate of volatiles even when this coat- Ehrenfreund P, Charnley SB (2000) Organic molecules in
ing comprises up to 10 % by mass of the starting the interstellar medium, comets, and meteorites: a
material (Gilmour et al. 2002). This implies that voyage from dark clouds to the early Earth. Ann Rev
the organic coating formed as one of the initial Astron Astrophys 38:427–483
Fegley B (1999) Chemical and physical processing of
products on any grain surface may also act as an presolar materials in the Solar Nebula and the impli-
additional effective catalytic surface for the con- cations for preservation of presolar materials in
version of H2, N2, and CO into organic materials. comets. Space Sci Rev 90:239–252
Fitness 857
Ferrante RF, Moore MH, Nuth JA, Smith T (2000) Labo- Definition
ratory studies of catalysis of CO to organics on grain
analogs. Icarus 145:297–300
Gilmour I, Hill HGM, Pearson VK, Sephton MA, Nuth JA Fitness is a central concept in evolutionary biol-
(2002) Production of high molecular weight organic ogy. It usually refers to the average capacity of an
compounds on the surfaces of amorphous iron silicate organism (as described by its ▶ genotype) to
catalysts: implications for organic synthesis in the produce viable progeny. The genotypes of fitter
Solar Nebula. LPSC 33:1613
Hayatsu R, Anders E (1981) Organic-compounds in mete- organisms become more abundant through the
orites and their origins. Top Curr Chem 99:1–37 action of ▶ natural selection along subsequent
Hill HGM, Nuth JA (2003) The catalytic potential of generations. Fitness is, however, a relative con-
cosmic dust: implications for prebiotic chemistry in cept since the same genotype, which is the unique
the solar nebula and other protoplanetary systems.
Astrobiology 3(2):291–304 heritable material, might express different ▶ phe-
Hindermann JP, Hutchings GJ, Kienneman A (1993) notypes in different environments. In turn, the F
Mechanistic aspects of the formation of hydrocar- ability of a phenotype (upon which natural selec-
bons and alcohols from CO hydrogenation. Catal Rev tion acts) to reproduce successfully depends on
Sci Eng 35:1–127
Hong Y, Fegley B (1998) Experimental studies of magne- the ▶ environment. The quantification of the rela-
tite formation in the Solar Nebula. Meteorit Planet Sci tionship between genotype and fitness remains an
33:1101–1112 open problem.
Johnson NM, Cody GD, Nuth JA (2004) Organics on
Fe-silicate grains: potential mimicry of meteoritic pro-
cesses? LPSC 35:1876
Johnson NM, Steiner ME, Nuth JA (2007) Fischer- Overview
Tropsch reactions and implications for protostellar
systems. LPSC 38:2183 Fitness is commonly defined as the ability of an
Kress ME, Tielens AGGM (2001) The role of Fischer-
Tropsch catalysis in Solar Nebula chemistry. Meteorit organism to pass its genes to the next generation.
Planet Sci 36:75–91 However, many different organismal and envi-
Llorca J, Casanova I (2000) Reaction between H2, CO, and ronmental traits are involved in the successful
H2S over Fe, Ni metal in the Solar Nebula: experimental completion of this process, and the mathematical
evidence for the formation of sulfur-bearing organic
molecules and sulfides. Meteorit Planet Sci 35:841–848 form of fitness is therefore difficult to cast. Often,
Nuth JA, Johnson NM, Manning S (2008) A self- the fitness of a single trait is instead addressed,
perpetuating catalyst for the production of complex that is, the advantages conferred by that trait to
organic molecules in protostellar nebulae. Astrophys the individuals carrying it. Actually, the first
J Lett 673(2):L225–L228
Pearce BB, Twigg MV, Woodward C (1989) In: Twigg quantification of fitness measured the relative
MW (ed) Catalyst handbook, 2nd edn. Wolfe, London, advantage of an organism with a mutant trait
pp 340–378 (or genotype) with respect to the non-mutated
Sabatier P, Senderens JB (1902) Direct hydrogenation of original type (Haldane 1924). Still, the fitness
oxides of carbon in presence of various finely divided
metals (in French). CR Acad Sci 134:689–691 value of a trait does not depend only on how it
performs in comparison to other genotypes, as the
case of the mutant allele causing sickle-cell ane-
Fitness mia illustrates. In a normal environment, homo-
zygous individuals may experience severe blood
Susanna Manrubia disorders. However, the disease confers certain
Systems Biology Program, Centro Nacional de resistance to malaria, enough for heterozygous
Biotecnologı́a (CSIC), Madrid, Spain individuals to be at an advantage in environments
where malaria is common, thus enhancing the
prevalence of sickle-cell anemia in those regions.
Keywords The definition of fitness as the number of
offspring of an organism has been challenged
Environment; Natural selection; Phenotype; since the beginning of evolutionary theory.
Reproduction; Survival Already Fisher discussed the case of two
858 Flint
individuals having the same number of offspring, Fisher RA (1930) The genetical theory of natural selec-
the first one breeding only female offspring and tion. Clarendon, Oxford
Haldane JBS (1924) A mathematical theory of natural and
the second one breeding half female and half artificial selection. Trans Camb Philos Soc 23:19–41
male offspring. Though they are equally fit in Hamilton WD (1964) The genetical evolution of social
terms of the definition above, the progeny of behaviour I and II. J Theor Biol 7:1-16–17-52
one of them cannot reproduce, and thus that lin- Orr EA (2009) Fitness and its role in evolutionary genet-
ics. Nat Rev Genet 10:531–539
eage would go extinct if left on its own (Fisher
1930). Hamilton extended the idea of fitness as
number of offspring (or number of genes that are
passed to the next generation) to include the
number of genes an individual can contribute to
Flint
the next generation by helping kin relatives to
▶ Chert
reproduce. This is called inclusive fitness
(Hamilton 1964).
Several issues have prevented the reaching of
a consensus definition for fitness. They include
whether fitness is better represented as a short-
Flood Basalt
term or as a long-term quantity; the role played by
▶ Trapps
chance and causality in the fate of populations;
whether it should be applied to species, organ-
isms, or genotypes; and how to consider its
dependence on the environment where organisms
reproduce and compete with each other (Byerly Flow Reactor
and Michod 1991; Abrams 2009). As a result,
fitness is used with different meanings in differ- Andrew Aubrey
ent situations, some of them being individual NASA Jet Propulsion Laboratory, Pasadena, CA,
fitness, absolute fitness, and relative fitness (Orr USA
2009).
Synonyms
See Also
Plug-flow reactor
▶ Adaptation
▶ Environment
Definition
▶ Genotype
A flow reactor is a temperature- and pressure-
▶ Natural Selection
controlled reaction vessel with opposite inlet
▶ Phenotype
and outlet ends for isothermal exposure of aque-
▶ Selection
ous or gaseous samples under laminar flow con-
▶ Survival
ditions. These systems consist of a narrow-bore
reactor with an in-line pump and back-pressure
References and Further Reading regulator and provide an alternative to batch-type
reactors. Residence time is equivalent to the ratio
Abrams M (2009) What determines biological fitness? of total reactor volume to the volumetric flow rate
The problem of the reference environment. Synthese
166:21–40
(tr ¼ V!), and variable exposures are achieved by
V
Byerly HC, Michod RE (1991) Fitness and evolutionary adjusting flow rate within the laminar flow
explanation. Biol Philos 6:1–22 regime. Flow reactors are commonly used to
Fluid Inclusions 859
Flow Reactor, Heating coil

Fig. 1 Temperature-
controlled flow reactor Effluent
system schematic
Feed Pump Flow reactor Cooling P-reg
solution (>100°C) stage
investigate aqueous chemical reactivity in high- Definition

temperature, high-pressure systems and are com-
monly utilized in astrobiology for investigation A fluid inclusion is a fluid phase present in the
into prebiotic organic synthetic pathways that rock porosity or filling fractures that is trapped in
might be possible at submarine hydrothermal microscopic cavities during mineral growth or F
systems (SHSs). High-temperature systems fracture healing. Inclusion fluids can be of sedi-
(100 C) include a rapid cooling stage before mentary, metamorphic, or magmatic origin. They
ambient sample collection (Fig. 1) to minimize can be thought of as time capsules storing infor-
hysteresis effects. Materials for high-temperature mation about ancient temperatures, pressures,
flow reactors include corrosion-resistant metals and fluid compositions relevant at the time of
such as high-nickel steel alloys or gold-plated trapping. The composition of most fluids falls in
surfaces. the C-O-H-S-N (H2O, CO2, CH4, H2S, N2) +
NaCl system, with NaCl representing the salt
content of the fluid (expressed in weight% NaCl
equivalent).
See Also
▶ Fischer-Tropsch-Type Reaction Overview

When crystals grow or recrystallize in a fluid
medium of any kind, growth irregularities of
many sorts trap microscopic portions of the ambi-
ent fluid in the solid crystal. These irregularities
are called fluid inclusions. The sealing off of such
Fluid Inclusions irregularities may occur during the growth of the
surrounding crystal, yielding primary fluid inclu-
Pascal Philippot sions, or by recrystallization along fractures at
Equipe Géobiosphère Actuelle et Primitive, some later time, yielding secondary inclusions.
Institut de Physique du Globe de Paris (IPGP), Fluid inclusions lining mineral growth zones are
Paris, France the best criteria for a primary origin. Secondary
inclusions occur generally along planes cutting
across several mineral grains. Fluid inclusions in
Keywords rock samples are seldom of a unique origin. The
presence of fluid inclusions in a rock is the rule
Fluid chemical composition; Fluid-rock interac- rather than the exception. Indeed, the rock sample
tion; Pressure; Homogenization temperature that contains no inclusions that are visible at high
magnification (>600) is rare, and many min-
erals such as milky quartz may contain up to 109
Synonyms fluid inclusions per cubic centimeter.
Fluid inclusions are present in all types of
Fluid-bearing micro cavities; Trapped fluids rocks, being sedimentary, metamorphic, or
860 Fluid Inclusions
igneous in origin. They have been traced down to phases at the time of trapping. Miscibility is a
mantle depths and were found in ultrahigh- function of both the salt content of the aqueous
pressure minerals such as diamond. They occur phase and the gas composition. One or more solid
in rocks as old as 4,000 Ma and have been iden- phases can also be present. Solid phases can be
tified in lunar and meteoric samples. Because either accidentally trapped particles that were
fluid inclusions are almost ubiquitous in geolog- present in suspension when the mineral grew or
ical samples, their study is applicable to a variety represent newly formed mineral that crystallized
of geological problems and areas. Fluid inclu- out of the fluids trapped in inclusions after
sions have played a central role in oil and ore sealing. In this case, the solid phase is called a
exploration research and in evaluating the safety daughter mineral.
of sites for both nuclear reactors and atomic Under the optical microscope, most fluid
waste repositories. Gas inclusions in polar ice inclusions have a rather sharp outer boundary
sheets have permitted reconstruction of CO2 con- marking the edge of the inclusion cavity. This is
centrations of the recent atmosphere, fluid inclu- because of a significant difference in refractive
sions in speleothems (cave deposits) have index between inclusion fluids and their mineral
provided data on paleotemperatures during the hosts: most aqueous fluids have refractive indices
last 350,000 years, and inclusions in samples between 1.33 and 1.45, whereas the minerals in
3,500 Ma old may provide geochemical con- which they are included have refractive indices
straints on the composition of the Earth’s primi- from 1.43 to as high as 3.22. Hydrocarbon liquids
tive atmosphere. In mantle and high-pressure have refractive indices that may be similar to
rocks, fluid inclusions have helped clarify the their mineral hosts and thus are not all easily
nature and role of volatiles in a variety of visible. Given appropriate fluid inclusions,
geodynamic processes including element many aspects of fluid composition can be deter-
recycling into the deep Earth, explosive volca- mined. The composition of most fluids falls in the
nism at convergent margins, and the seismic C-O-H-S (H2O, CO2, CH4, H2S) + NaCl system,
behavior of the subducting plate. In metamorphic with NaCl representing the salt content of the
and sedimentary terranes, fluid inclusions have fluid (expressed in weight% NaCl content) and
proven useful tools for containing the burial and accounting for KCl, CaCl2, and other chlorides.
exhumation history of crustal rocks and the pres- Some fluid inclusions can also contain significant
sure and temperature changes attending uplift and amounts of nitrogen (N2). Bromine and fluorine
erosion of continental masses. can be important volatiles in some systems, but
Fluid inclusions can be thought of as time their abundances (as HBr and HF) are generally
capsules storing information about ancient tem- very low. In sedimentary rocks, oil inclusions can
peratures, pressures, and fluid compositions rele- be present. The nature and concentrations of a
vant at the time of trapping. The basic assumption large variety of major and trace elements can be
in all fluid inclusion studies is that the volume obtained by crushing and leaching the inclusion
(density) and composition of the trapped fluid has fluids out of small samples containing a large
not changed since formation of the inclusion or number of inclusions or in situ, within a single
that, if an inclusion did leak, evidence for fluid inclusion. Information on the isotopic composi-
loss is observable in the sample (decrepitation tion of light elements (H, C, N, O, S, and some
texture, microfracture, etc.). With the exception noble gases) of the fluid can also be obtained.
of magmatic rocks that can contain glass inclu- If the composition of the inclusion can be
sions, that is, solidified melt inclusions, most determined and the liquid-vapor curve for that
rocks contain liquid and/or gas inclusions. At fluid is known, then the density can be deter-
room temperature, a general distinction can be mined from the temperature of homogenization
made between liquid and gaseous inclusions. of the vapor and liquid phases to one fluid phase.
Both types are often found in the same rock. If the P-V-T properties of that fluid are known,
This might indicate immiscibility between two then a line of constant density or molar volume is
Fluid Inclusions 861
defined in a P-T space. This line, called an inclusions and opened the possibility for the
isochore, represents the range of P-T conditions determination of fluid salinities and densities,
over which a fluid of that density was trapped. If and the consequent interpretation on the rock-
one can make an independent estimate of pres- forming conditions. Numerous analytical
sure or temperature at the time of trapping of the methods were developed since then for measur-
fluid in an inclusion, then both variables (P and T) ing fluid chemical and isotopic compositions.
can be uniquely determined using the isochore of Among these, the introduction of laser Raman
the fluid. If both aqueous and gaseous inclusions spectroscopy around 1980 (Dubessy et al. 1982;
are coexisting in the same healed fracture or fluid Pasteris et al. 1988) and synchrotron radiation
inclusion cluster, and the specific criteria for micro-x-ray fluorescence (Frantz et al. 1988;
immiscibility between these two phases at the Vanko et al. 1993; Philippot et al. 1998), Laser
time of trapping can be ascertained precisely, Ablation Inductively Coupled Plasma Mass F
then the intersection point between the two Spectrometry (Heinrich et al. 2003), and Laser
isochores in a P-T space corresponds to the tem- Microprobe Noble Gas Mass Spectrometry
perature and pressure of trapping of the inclusion (Bohlke and Irwin 1992) around 1990 allowed
fluids. quantitative analysis of anhydrous gas species
(CO2, N2, CH4, H2S, and from then on H2O),
major and trace elements, as well as noble gases
History and Basic Methodology in individual fluid inclusions. Much of this work
has been reviewed in the treatise on fluid inclu-
The presence of fluids in rocks has been noticed sions published by Roedder (1984). Some of the
since at least the eleventh century, but the first important concepts and technical achievements
fluid inclusion constituents (H2O, CO2, petro- performed during this period are synthesized in
leum) were identified not before the eighteenth Hollister and Crawford (1981) and Goldstein and
and nineteenth centuries (Dolomieu 1792; Sorby Reynolds (1994).
1858). Sorby (1858) was the first to highlight a
relation between metamorphism and filling of Fluid Inclusions in Meteorites and Early
fluid inclusions. He suggested that the gas-liquid Archean Sediments
phases in fluid inclusions reflected thermally The discovery of water-bearing fluid inclusions
contracted fluids. He proposed that reheating in 4.5 billion-year-old halite (NaCl) crystals pre-
inclusions would lead to disappearance of the served in the matrix of the Monahans and Zag
bubble and that the temperature at which this H-▶ chondrite regolith ▶ breccias (Zolensky
occurred could serve as an estimate of the tem- et al. 1999) provided important constraints on
perature of mineral formation. During the twen- the fluid composition and P-T conditions in the
tieth century, fluid inclusions were the subject of near-surface environment of the ▶ parent bodies.
intense studies and debates by geologists from Minimum formation temperature of the inclu-
Russia, the USA, and Europe. The development sions has been estimated to be less than 100 C
and improvement of the equipment for quantita- and could have been as low as about 0.01 C. The
tive analysis of fluid inclusions during the late minimum pressure needed to stabilize a NaCl-
1960s and early 1970s paved the way to a vast saturated brine is 0.0754 MPa at 100 C and
number of geological applications using fluid 3 104 MPa at 0.01 C. These conditions are
inclusions as a petrological tool. Especially, a consistent with the alteration having occurred in
cooling-heating stage was developed (Poty the shallow subsurface of the parent bodies.
et al. 1976), which expanded to low temperatures Moreover, the P-T conditions inferred for the
the temperature range for easily and accurately formation of these inclusions is within the range
measuring phase transition in fluid inclusions. of temperatures in which life has been observed
This technique known as “microthermometry” to exist and thrive on Earth. Tiny fluid inclusions
is indispensable for the analysis of single fluid have also been found in two Martian meteorites
862 Fluid Inclusions
(one in ▶ Nakhla, NSNM 5891–3, and one in was thought to represent a relic testimony of the
▶ ALH 84001, 146) (Bodnar 2000) and carbona- primitive ocean. By contrast, Ba- and Fe-rich
ceous chondrites (Saylor et al. 2001). Although brines have Cl/Br ratios (350 and 390) close to
the fluid inclusions in the Nakhla sample occur as bulk Earth value (420), hence arguing for a man-
healed fracture and therefore are of secondary tle buffering and a hydrothermal origin of these
origin, the inclusions in the ALH 84001, 146 sam- fluids. Model composition of “seawater compo-
ple do not occur along fractures and therefore nent” has been evaluated to be 1,100 mM Na,
could be of primary origin, that is, trapped during 2,250 mM Cl, and 375 mM Ca, which corre-
growth of the enclosing pyroxene. The fluid in the sponds to a bulk fluid salinity of 12 wt% salt
inclusions may represent the magmatic fluid that equivalent. This high Cl concentration (ca. four
exsolved from the crystallizing melt as the igne- times the present-day value) together with the
ous rock formed on Mars. As such, the inclusions occurrence of Cl/Br ratio similar to the modern
can provide valuable information about seawater value (649) indicates that the Archean
degassing early in Mars history. Optical behavior ocean formed on a typical modern-day seawater
of these inclusions suggests that they contain evaporation trend.
liquid and vapor carbon dioxide.
Insights into the composition of the primitive
terrestrial ocean and hydrothermal fluids (Cl/Br See Also
ratio, sulfur, and metal components) have been
obtained by direct analysis of fluid inclusions ▶ Archean Eon
preserved in poorly metamorphosed Archean ▶ Archean Traces of Life
sediments of the Kaapvaal Craton (South Africa) ▶ Geothermobarometers
and Pilbara Craton (Western Australia). Early ▶ North Pole Dome (Pilbara, Western Australia)
studies performed in 3.2 Gyr-old (Channer ▶ Ocean, Chemical Evolution of
et al. 1997; De Ronde et al. 1997) (but see Lowe ▶ Syngenicity
and Byerly (2003) for a controversy on the origin
of the samples considered) and paleoproterozoic
2.2 Gyr-old (Gutzmer et al. 2003) indicated that References and Further Reading
the Cl/Br ratio of the fluid trapped in the inclu-
Bodnar RJ (1983) A method of calculating fluid inclusion
sions was below present-day seawater and volumes based on vapor bubble diameters and P-V-T-
resulted from mantle buffering, that is, hydrother- X properties of inclusion fluids. Econ Geol
mal in origin. However, these studies were based 78:535–542
on bulk fluid analyses (i.e., crush-leach), which Bodnar RJ (2000) Fluid inclusions in ALH 84001 and
other Martian meteorites: evidence for volatiles on
can result in fluid mixing if several fluid genera- Mars. Lunar Planet Sci 30:1222
tions occur in a single sample. More recently, Bohlke JK, Irwin JJ (1992) Laser microprobe analyses
chemical analysis of individual fluid inclusions of noble gas isotopes and halogens in fluid
have been performed in 3.5 Gyr-old samples inclusions: analyses of microstandards and synthetic
inclusions in quartz. Geochim Cosmochim Acta
from the Pilbara Craton, Western Australia, by 56:187–201
Synchrotron Radiation X-ray micro- de Channer DMR, de Ronde CEJ et al (1997) The Cl, Br,
Fluorescence (SR-XRF), thus allowing indepen- I composition of f3.23 Ga modified seawater: implica-
dent analysis of different fluids trapped in the tions for the geological evolution of ocean halide
chemistry. Earth Planet Sci Lett 150:325–335
same sample (Foriel et al. 2004). Individual de Dolomieu CD (1792) Sur l’huile de pétrole dans le
fluid inclusion analysis using SR-XRF revealed cristal de roche et les fluides élastiques tirés du quartz.
the presence of three main fluid populations: a Obs Phys 42:318–319
metal-depleted fluid, a Ba-rich and S-depleted De Ronde EJ, Channer DMD, Faure K, Bray CJ, Spooner
ETC (1997) Fluid chemistry of Archean seafloor
fluid, and a Fe-S-rich end-member. The Cl/Br hydrothermal vents: implications for the composition
ratio of metal-depleted fluid inclusions (630) is of circa 3.2 Ga seawater. Geochim Cosmochim Acta
similar to the modern seawater value (649) and 61:4025–4042
Fluorescence 863
Dubessy J, Audeoud D, Winkins R, Kosztolanyi C (1982) Vanko DA, Sutton SR, Rivers ML, Bodnar RJ
The use of the Raman microprobe MOLE in the deter- (1993) Major-element ratios in synthetic fluid inclu-
mination of the electrolytes dissolved in the aqueous sions by synchrotron x-ray fluorescence microprobe.
phase of fluid inclusions. Chem Geol 37:137–150 Chem Geol 109:125–134
Foriel J, Philippot P, Rey P, Somogyi A, Banks D, Ménez Zolensky ME, Bodnar RJ, Gibson EK Jr, Nyquist LE,
B (2004) Biological control of Cl/Br and low sulfate Reese Y, Shih CY, Wiesmann H (1999) Asteroidal
concentration in a 3.5-Gyr-old seawater from North water within fluid inclusion-bearing halite in an H5
Pole, Western Australia. Earth Planet Sci Lett chondrite, Monahans (1998). Science 285:1377–1379
228:451–463
Frantz JD, Mao HK, Zhang YG, Wu Y, Thompson AC,
Underwood JH, Giauque RD, Jones KW, Rivers ML
(1988) Analysis of fluid inclusions by x-ray fluores-
cence using synchrotron radiation. Chem Geol Fluid-Bearing Micro Cavities
69:235–244
Goldstein RH, Reynolds TJ (1994) Systematics of fluid
▶ Fluid Inclusions F
inclusions in diagenetic minerals, vol 31, Society for
sedimentary geology, short course. SEPM, Tulsa,
199 pp
Gutzmer J, Banks DA, Luders V, Hoefs J, Beukes NJ, Von
Bezing KL (2003) Ancient sub-seafloor alteration of
basaltic andesites of the Ongeluk Formation, South
Fluorescence
Africa: implications for the chemistry of Paleopro-
terozoic seawater. Chem Geol 201:37–53 Henderson James (Jim) Cleaves II
Heinrich CA, Pettke T, Halter WE, Aigner-torres M, Earth–Life Science Institute (ELSI), Tokyo
Audétat A, Gunther D, Hattendorf B (2003) Quantita-
tive multi-element analysis of minerals, fluid and melt
inclusions by laser-ablation inductively coupled- Japan
plasma mass-spectrometry. Geochim Cosmochim Institute for Advanced Study, Princeton, NJ,
Acta 67(18):3473–3496 USA
Hollister LS, Crawford ML (1981) Short course in
fluid inclusions: applications to petrology. Mineral
Association of Canada, Calgary, Special Publication, Washington, DC, USA
304 pp Center for Chemical Evolution, Georgia Institute
Lowe DR, Byerly GR (2003) Ironstone pods in the of Technology, Atlanta, GA, USA
Archean Barberton Greenstone Belt, South Africa:
Earth’s oldest seafloor hydrothermal vents
reinterpreted as Quaternary subaerial springs. Geology
31:909–912 Synonyms
Pasteris JD, Wopenka B, Seitz JC (1988) Practical aspects
of quantitative laser Raman microprobe spectroscopy
Fluorescent emission
for the study of fluid inclusions. Geochim Cosmochim
Acta 52:979–988
Philippot P, Ménez B, Chevalier P, Gibert F, Legrand F,
Populus P (1998) Absorption correction procedures for Definition
quantitative analysis of fluid inclusions using synchro-
tron radiation x-ray fluorescence. Chem Geol
144:121–136 When an electron in a substance is excited to a
Poty B, Leroy J, Weisbrod A (1976) A new device for higher energy state by a photon, it may return to
measuring temperatures under the microscope: the the lower energy state via series of two or more
Chaixmeca microthermometry apparatus. Bull Soc Fr
intermediate energy states, assuming they exist.
Minéral Cristallogr 99:182–186
Roedder E (1984) Fluid inclusions. Mineralogical Society The photons which are re-emitted have a lower
of America, Washington, DC, 644 pp energy than the exciting photon and are conse-
Saylor J, Zolensky M, Bodnar R, Le L, Schwandt C (2001) quently of a lower frequency and longer wave-
Fluid inclusions in carbonaceous chondrites. Lunar
Planet Sci 32:1885
length. This phenomenon is known as
Sorby HC (1858) On the microscopic structure of crystals, fluorescence. In some instances, two electrons
indicating the origin of minerals and rocks. Q J Geol may be adsorbed, allowing for emission of radi-
Soc Lond 14:453–500 ation of a shorter wavelength. As in UV
864 Fluorescence Resonance Energy Transfer
spectroscopy, compounds often have unique fluo-

rescence spectra which can be used in determin- Fluorometry
ing structure (see ▶ Fluorometry). Fluorescence
detectors are extremely sensitive, thus this is an Michael P. Callahan
excellent analytical technique for trace amounts Astrochemistry Laboratory, Code 691, NASA
of substances. The infrared emission bands Goddard Space Flight Center, Greenbelt, MD,
assigned to ▶ polycyclic aromatic hydrocarbons USA
(PAHs) in a variety of astronomical environ-
ments have been attributed to fluorescence, with
the exciting radiation being in the ultraviolet
Synonyms
region of the spectrum.
Fluorescence spectroscopy; Fluorimetry;
Spectrofluorometry
See Also

Definition
Fluorometry is a chemical technique for

measuring or monitoring ▶ fluorescence. In fluo-
Fluorescence Resonance Energy rometry, the species is excited from its
Transfer ground electronic state to an excited electronic
state by absorption of a ▶ photon (typically from
▶ FRET a UV source). The species returns to the
ground electronic state by emitting one
(resonance fluorescence) or more photons.
The amount of fluorescence measured is
Fluorescence Spectroscopy used to determine the sample concentration by
comparison with a standard or by using a calibra-
▶ Fluorometry tion curve. Fluorometry is highly sensitive with
up to femtomolar detection limits. In addition,
fluorometry is considered a more specific tech-
nique than ▶ absorption spectroscopy, since fluo-
rescent molecules (fluorophores) are less
Fluorescent Emission common.
▶ Fluorescence
See Also
▶ Absorption Spectroscopy
Fluorimetry ▶ Fluorescence
▶ Fluorophore
▶ Fluorometry ▶ Photon
Fomalhaut b 865
Fluorophore Flux, Radiative
Henderson James (Jim) Cleaves II Daniel Rouan

Earth–Life Science Institute (ELSI), Tokyo LESIA, Observatoire Paris-Site de Meudon,
Institute of Technology, Meguro–ku, Tokyo, Japan Meudon, France
Washington, DC, USA Definition
of Technology, Atlanta, GA, USA The radiative flux is the amount of electromag-
netic power crossing one surface unit. In astron- F
omy, it corresponds to the part of the power
Keywords emitted by a celestial object received per area
unit at the Earth. If the contributions at all the
Chromophore wavelengths are summed, one speaks of bolomet-
ric flux and the unit is W m2. If only a small
interval in frequency (or wavelength) is consid-
ered, one speaks of spectral flux (unit: W m2
Definition Hz1). In the radio domain, the unit currently
used is the Jansky (1026 W m2 Hz1).
A fluorophore is a molecule or a portion of a
molecule that has fluorescent properties. While
many compounds are inherently fluorescent as See Also
they contain fluorophores, there are many com-
pounds that are not. For the sake of analyzing ▶ Electromagnetic Radiation
these latter types of molecules, a fluorophore is ▶ Magnitude
often introduced via chemical derivatization
with a fluorescent reagent, for example, fluores-
cein isothiocyanate (FITC) or fluorescamine.
Many widely used DNA stains and cell stains
such as 40 , 6-diamidino-2-phenylindole (DAPI) Fomalhaut b
are fluorophore-containing molecules.
In the interstellar medium, infrared emission Marc Kuchner1 and Nader Haghighipour2
1
attributed to ▶ polycyclic aromatic hydrocarbon NASA Goddard Space Flight Center,
molecules (PAHs) is thought to result from fluo- Exoplanets and Stellar Astrophysics Laboratory,
rescence following the absorption of ultraviolet Greenbelt, MD, USA
2
photons. Institute for Astronomy, University of Hawaii-
See Also
Synonyms
▶ Fluorometry
▶ Polycyclic Aromatic Hydrocarbon a Piscis Austrinus b
866 Fomalhaut b
Fomalhaut b,
Fig. 1 False-color,
unsmoothed image of the
2012 STIS data. The box
inset is 1 . on a side and
magnifies the image of
Fomalhaut b. North is up,
east is left (Image from
Kalas et al. 2013)
Fomalhaut b, Fig. 2 Northwest portions of the 2012 both data sets. Ovals indicate background galaxies and
data (left) and 2010 data (right) with 5 5 median binning Fomalhaut b is marked between white line segments
and a hard grayscale stretch to emphasize the northwest (Image from Kalas et al. 2013)
gap. The dotted lines are at P.A. = 329.8 . and 332.8 in
Definition main sequence A-type star Fomalhaut, 25 light

years from Earth. Paul Kalas, James Graham, and
Fomalhaut b is a massive planet orbiting roughly Mark Clampin discovered the planet in 2008 in a
115 AU (▶ AU = astronomical units) from the series of images of the star made by the ▶ Hubble
Formaldehyde 867
Space Telescope (HST), showing the object to

orbit the star just inside a narrow ring of dust. Formaldehyde
The planet’s mass is constrained to the
range 0.05–3 Jupiter masses based on the shape Henderson James (Jim) Cleaves II
of the dust ring, which the planet stirs gravita- Earth–Life Science Institute (ELSI), Tokyo
tionally. The ring of dust around Fomalhaut Institute of Technology, Meguro–ku, Tokyo, Japan
tipped off observers in the 1980s that this Institute for Advanced Study, Princeton, NJ, USA
star might host a planetary system. Images of Blue Marble Space Institute of Science,
this narrow ring with the HST showed Washington, DC, USA
that the ring is not centered on the star; its center Center for Chemical Evolution, Georgia Institute
is offset from the star by 15 AU. This offset of Technology, Atlanta, GA, USA
suggested the presence of a planet perturbing F
the ring, and led to the planet’s discovery in
2008. The planet remains mysterious; it is several Keywords
times brighter than expected for a body with this
mass, and it has so far escaped observers who Formose; Prebiotic chemistry; Strecker synthesis
have tried to image it using other instruments.
Fomalhaut b is considered the first
extrasolar planet directly imaged by the HST; it Synonyms
is roughly a billion times fainter than the star it
orbits (Figs.1 and 2). Methanal; Methyl aldehyde; Methylene oxide;
Paraformaldehyde
See Also Definition
▶ Beta Pictoris b Formaldehyde (methanal, HCHO), the simplest

▶ Direct-Imaging, Planets ▶ aldehyde, is a one carbon molecule intermedi-
▶ Exoplanets, Discovery ary along the redox continuum between CO2 and
▶ HR 8799: The First Directly Imaged Multi- CH4, at the same oxidation state (0) as ▶ graphite.
Planet System Formaldehyde was first reported by the Russian
chemist Butlerow in 1860 and was conclusively
identified by von Hofmann. It exists transiently but
prominently in the abiological carbon cycle
Kalas P, Graham JR, Chiang E, Fitzgerald MP, (Cleaves 2008). It is an abundant interstellar mol-
Clampin M, Kite ES, Stapelfeldt K, Marois C, Krist ecule and is a constituent of cometary ices (Biver
J (2008) Optical images of an exosolar planet 25 light- et al. 2002). It is readily produced in prebiotic
years from Earth. Science 322(5906):1345
simulation experiments from a variety of gas mix-
Kalas P, Graham JR, Fitzgerald MP, Clampin M (2013)
STIS coronagraphic imaging of Fomalhaut: main belt tures and energy sources.
structure and the orbit of Fomalhaut b. The Astrophys HCHO may have played an important role in
J 775(1):31, Article id 56 the synthesis of organic molecules relevant to the
origin of life. HCHO is likely a significant pre-
cursor for the prebiotic synthesis of ▶ glycine and
other amino acids (Miller 1957). HCHO can react
to form sugars under basic conditions (Butlerow
Footballene 1861) via the so-called ▶ formose reaction, a
reaction of potential importance for the origin of
▶ Fullerene an “▶ RNA World” or early nucleic acids based
868 Formaldehyde
on alternative sugars in a potential “pre-RNA (see below) (Calvert and Steacie 1951). Sekine
World” (Joyce et al. 1987). It has been argued (2002) found that MnO2 catalyzes the oxidation
that HCHO may in fact be the only one carbon C-, of HCHO at room temperature to CO2 under
H-, and O-containing molecule capable of gener- atmospheric conditions. Löb (1906) found that
ating complex organic compounds for the origin electric discharges acting on HCHO in the pres-
of life (Weber 2002). ence of water vapor give CO, CO2, H2, and CH4
as products, which is essentially the reverse of the
reaction demonstrated by Miller (1953).
Overview There is reason to believe that there was a
considerable flux of atmospherically synthesized
Prebiotic Sources of HCHO and rained out H2O2 to the primitive oceans,
One important source of HCHO on the primitive particularly if the atmosphere was nonreducing
Earth is atmospheric photochemical synthesis by (Kasting and Brown 1998). Reaction of HCHO
the photoreduction of CO2 with H2O: with aqueous hydrogen peroxide gives formate
and H2 (Walker 1964) (Reaction 3):
CO2 þ H2 O $ HCHO þ O2 (1)
2HCHOðaqÞ þ 2OH þ H2 O2 ðaqÞ
Pinto et al. (1980) estimated that HCHO could be $ 2HCOO ðaqÞ þ 2H2 O þ H2 ðgÞ (3)
produced in yields of up to 1011 moles/year,
reaching a steady-state oceanic concentration of Prebiotic Solution Chemistry of HCHO
103 M in 107 years. Electric discharges acting on An excellent monograph has been written on the
a variety of gas mixtures also produce HCHO in chemistry of HCHO (Walker 1964). In aqueous
good yield (Stribling and Miller 1987); however, solution, HCHO is mostly present as the
photochemistry may have been more significant monohydrate, methylene glycol (Eq. 4):
simply due to the greater energy flux. Photochem-
ical production from atmospheric CH4 and/or CO HCHO þ H2 O $ CH2 ðOHÞ2 (4)
could also have been a significant source of
HCHO, though the production and rainout rates HCHO absorbs UV radiation strongly in the range
of HCHO are highly sensitive to energy source and of 250–350 nm, resulting in photolysis. The
atmospheric composition (Chang 1993). hydrate does not absorb light at these wavelengths,
HCHO could also potentially be produced in thus dissolution in water could be an important
hydrothermal vents (Ferris 1994), for example, by photoprotection mechanism for HCHO.
the reduction of aqueous formate, CO or CO2, or by
the oxidation of methanol or CH4, although HCHO
Oligomerization Chemistry:
does not appear to be a stable redox state of carbon
Polyoxymethylene
under hydrothermal conditions. HCHO may be an
Low molecular weight polymers,
intermediate in the redox equilibration of other
▶ polyoxymethylene (POMs) (of formula HO
more stable compounds (Seewald et al. 2006).
(CH2O)nH), form readily in neutral concentrated
aqueous HCHO solutions. The cyclic oligomers,
Prebiotic Sinks for HCHO
trioxane and tetraoxane (Fig. 1), may also form
HCHO decomposes thermally in the gas phase
where acid catalysis is available.
above 300 C to give CO and H2 (Bone and Smith
1905) (Eq. 2):
Oligomerization Chemistry: The Formose
HCHOðgÞ $ COðgÞ þ H2 ðgÞ (2) Reaction
HCHO can be oligomerized under basic condi-
Long wavelength photochemical destruction tions into sugars via the so-called formose reac-
was also likely a significant sink for HCHO tion (Butlerow 1861) (see ▶ Formose Reaction).
Formaldehyde 869
The equilibrium constant for the dimerization of et al. 1980). If HCHO became sufficiently con-
HCHO to give ▶ glycolaldehyde is not well centrated, side reactions may have established the
known, but is of considerable interest, since steady-state concentration regardless of the pro-
while HCHO is volatile, glycolaldehyde is not duction rate. Given the observed photochemical
and could thus be concentrated by evaporation. synthesis of pentaerythritol (Fig. 2, left hand side)
HCHO reacts with ▶ acetaldehyde at much (Schwartz and De Graaf 1993), and its likely
lower concentrations to give acrolein among mechanism of synthesis, it seems possible that
other products (Cleaves 2003). HCHO is converted to CH3CHO by UV light,
The reaction of HCHO into formose is one and thus it appears unlikely that bulk oceanic
limiting reaction pathway for the accumulation concentrations of HCHO could have been much
of HCHO in primitive waters, which would higher than 103 M.
depend on the rate of HCHO production and HCHO is prone to photoreaction in the primi- F
delivery to the primitive oceans (Pinto tive oceans. For example, HCHO photochemically
reacts to give mostly pentaerythritol (Schwartz
O and De Graaf 1993), among other nonsugar prod-
O ucts (Fig. 2) (Shigemasa et al. 1977).
O
Reactions with Amines
HCHO is converted to ▶ hexamethylenetetra-
mine (HMT or 1, 3, 5, 7-tetraazatricyclo
O O O
[3.3.1.13,7] decane) (Fig. 3) by reaction with
ammonia (NH3).
O
The reaction of fairly dilute ( 0.003 M) aque-
Formaldehyde, Fig. 1 Cyclic oligomers of HCHO, ous HCHO and NH3 between 100 C and 250 C
trioxane, and tetraoxane produces amino acids, amines, and amino
Formaldehyde, Fig. 2 2- HO HO OH
Hydroxymethylglycerol
and pentaerythritol: major
products of photochemical
reactions of HCHO OH
HO HO
HO HO
Formaldehyde, Fig. 3 N
Hexamethylenetetramine
(HMT) forms readily from
NH3 and HCHO
6 HCHO + 4 NH3 N N + 6 H2O
Hexamethylenetetramine
870 Formaldehyde
not especially high. This is generally also true for

low molecular weight POMs (Parfitt and Green-
S S land 1970). The adsorption equilibria may be
more significant for clays such as illite and
▶ kaolinite (Chandra and De 1983). Clay adsorp-
tion may have led to significant concentration
effects which may have facilitated reactions of
HCHO such as oligomerization to formose prod-
S ucts and redox reactions to ▶ methanol, formate,
Trithiane and CO2.
Formaldehyde, Fig. 4 Trithiane is formed from H2S Reaction with HCN

and HCHO Aqueous HCHO reacts readily with HCN to give
glycolonitrile (Henry 1890) (Eq. 9):
alcohols (Aubrey et al. 2009), and the reaction of
somewhat more concentrated HCHO (0.02 M) HCHO þ HCN $ HOCH2 CN (9)
with NH3 (0.04 M) has been shown to produce
traces of amino acids at lower temperatures The Keq for the reaction is exceptionally high
(40 C) (Weber 1998). Thus, for at least several (Schlesinger and Miller 1973); thus, any atmo-
simple amino acids, NH3 and HCHO may be spheric composition which results in the produc-
sufficient for synthesis to occur. tion of the two compounds essentially results in
the production of ▶ glycolic acid, after hydroly-
Reactions with Sulfur Species sis of the ▶ nitrile.
HCHO and H2S react rapidly to produce trithiane (Schwartz and Goverde 1982) found that the
and other oligomers (Fig. 4), which have been addition of HCHO or glycolonitrile to HCN actu-
detected in hydrothermal vent effluent (Simoneit ally accelerates the formation of HCN tetramer
1992), as well as in hydrothermal vent simula- diaminomaleonitrile (DAMN). HCHO reacts
tions (Cole et al. 1994) (Eqs. 5, 6, and 7): readily with DAMN to form a crystallizable prod-
uct (Koch et al. 2007).
H2 S þ H2 CðOHÞ2 $ HOCH2 SH þ H2 O (5) Reaction of HCN, HCHO, and NH3 produces
the amino acid glycine via the Strecker synthesis.
2 HOCH2 SH $ HSCH2 OCH2 SH (6) Cannizzaro reactions may also be important
loss channels for HCHO. If HCHO concentra-
HOCH2 SH þ HSCH2 OCH2 SH tions are high enough, HCHO disproportionates
$ HSCH2 SCH2 OCH2 SH þ H2 O (7) to form formate and methanol (Walker 1964),
which reduces the concentration of HCHO avail-
Aqueous HCHO reacts with SO2 to give able to form sugars. These reactions are acid,
methylolsulfonic acid (Walker 1964) (Eq. 8): base, and metal catalyzed (Walker 1964). As the
reaction is apparently third order, it is unclear
H2 CO þ H2 O þ SO2 $ HOCH2 SO3 H (8) whether it would occur significantly in dilute
solution.
The salts of this compound are nonvolatile, but
decompose in dilute acid, liberating HCHO. Interstellar Formaldehyde
Formaldehyde was the first polyatomic organic
Mineral Interactions molecule detected in the interstellar medium by
There have been few studies of the adsorption of Zuckerman et al. (1970). Since its initial detec-
HCHO to minerals, though the ones that do exist tion, it has been observed in many regions of the
suggest that HCHO clay adsorption equilibria are galaxy. As the gas-phase reaction that produces
Formaldehyde 871
formaldehyde is too inefficient to produce the Calvert A, Steacie E (1951) The vapor phase photolysis of
abundance of formaldehyde that has been formaldehyde at wavelength 3130. J Chem Phys
19:1976–1982
observed, it has been proposed that HCHO is Chandra K, De S (1983) Adsorption of formaldehyde by
formed via the hydrogenation of CO in ice clay minerals in presence of urea and ammonium sul-
(Eqs. 10–11): fate in aqueous system. Indian J Agric Chem
16:239–245
Chang S (1993) Prebiotic synthesis in planetary environ-
H þ CO ! HCO (10) ments. In: Greenberg JM, Mendoza-Gomez CX,
Pirronello V (eds) The chemistry of life’s origins.
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Cleaves H (2003) The prebiotic synthesis of acrolein.
Monatsh Chem 134:585–593
There are several side reactions that take place Cleaves HJ (2008) The prebiotic geochemistry of formal-
with each step of the reaction that depend on the dehyde. Precambrian Res 164(3–4):111–118 F
nature of the ice on the grain (Woon 2002). The Cole W, Kaschke M, Sherringham J, Curry G, Turner D,
isotopic ratio of [12C]/[13C] in HCHO in the Russell M (1994) Can amino acids be synthesized by
H2S in anoxic lakes? Mar Chem 45:243–256
galactic disk has been determined to be 50:1 Ferris J (1994) The potential for prebiotic synthesis in
(Henkel et al. 1985), while the terrestrial ratio is hydrothermal systems. Orig Life Evol Biosph
100:1. 24:363–381
Fox S, Windsor C (1970) Synthesis of amino acids by the
heating of formaldehyde and ammonia. Science
Formaldehyde in Biology 170:984–986
Free formaldehyde is rarely found in living Gabel N, Ponnamperuma C (1967) Model for origin of
metabolism, as formaldehyde is mutagenic due monosaccharides. Nature 216:453–455
to its reactivity with proteins and nucleic acids. Henkel C, Guesten R, Gardner FF (1985) [12C]/[13C]
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tetrahydrofolate; however, in methanogenesis, Henry L (1890) Sur le nitrile gycolique et la synthèse
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Joyce G, Schwartz A, Miller S, Orgel L (1987) The case
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logues of the nucleotides. Proc Natl Acad Sci U S
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atmosphere as a source of biogenic compounds. In:
▶ Aldehyde Brack A (ed) The molecular origins of life: assembling
▶ Formose Reaction the pieces of the puzzle. Cambridge University Press,
New York, pp 35–56
▶ Strecker Synthesis Koch K, Schweizer W, Eschenmoser A (2007) Reactions
of the HCN-tetramer with aldehydes. Chem Biodivers
4:541–553
References and Further Reading Löb W (1906) Studien € uber die chemische Wirkung der
stillen elektrischen Entladung. Zeitschr f€ ur
Aubrey AD, Cleaves HJ, Bada JL (2009) The role of Elektrochem 15:282–312
submarine hydrothermal systems in the synthesis of Miller S (1953) A production of amino acids under
amino acids. Orig Life Evol Biosph 39(2):91–108 possible primitive Earth conditions. Science
Biver N, Bockelée-Morvan D, Crovisier J, Colom P, 117:528–529
Henry F, Moreno R, Paubert G, Despois D, Lis Miller SL (1957) The mechanism of synthesis of amino
D (2002) Chemical composition diversity among acids by electric discharges. Biochim Biophys Acta
24 comets observed at radio wavelengths. Earth 23(3):480–489
Moon Planet 90:323–333 Parfitt R, Greenland D (1970) The adsorption of poly
Bone W, Smith H (1905) The thermal decomposition of (ethylene glycols) on clay minerals. Clay Miner
formaldehyde and acetaldehyde. J Am Chem Soc 8:305–315
87:910–916 Pinto J, Gladstone G, Yung Y (1980) Photochemical pro-
Butlerow A (1861) Formation synthétique d’une subduction of formaldehyde in Earth’s primitive atmo-
stance sucrée. Compt Rendus Acad Sci 53:145–147 sphere. Science 210:183–185
872 Formamide
Schlesinger G, Miller S (1973) Equilibrium and kinetics of Definition

glyconitrile formation in aqueous solution. J Am Chem
Soc 95:3729–3735
Schutte W, Allamandola L, Sandford S (1993) An exper- Formamide (IUPAC name, methanamide) is the
imental study of the organic molecules produced in ▶ amide with the simplest structure (other
cometary and interstellar ice analogs by thermal form- amides have one, two, or three hydrogen atoms
aldehyde reactions. Icarus 104:118–137 replaced by radicals). Formamide is the smallest
Schwartz A, De Graaf R (1993) The prebiotic synthesis of
carbohydrates: a reassessment. J Mol Evol 36:101–106 molecule with a peptide bond. At room tempera-
Schwartz A, Goverde M (1982) Acceleration of HCN ture and standard pressure, it is a colorless liquid.
oligomerization by formaldehyde and related com- It remains liquid between 3 and 210 C: this large
pounds: implications for prebiotic syntheses. J Mol range is one of the reasons for which formamide
Evol 18:351–353
Seewald JS, Zolotov M, McCollom T (2006) Experimental was proposed as an alternative to water as the
investigation of single carbon compounds under hydrother- “ideal” solvent for life. Above 90 C, it decom-
mal conditions. Geochim Cosmochim Acta 70:446–460 poses to HCN and water. Ammonium formate,
Sekine Y (2002) Oxidative decomposition of formalde- itself a product of the reaction of the two simple
hyde by metal oxides at room temperature. Atmos
Environ 36:5543–5547 species, formic acid and ammonia, produces
Shigemasa Y, Matsuda Y, Sakazawa C, Matsuura formamide when heated. Formamide is a poten-
T (1977) Formose reactions II. The photochemical tial starting material to create the RNA bases
formose reaction. Bull Chem Soc Jpn 50:222–226 (Barks et al. 2010). Formamide has been found
Simoneit B (1992) Aqueous organic geochemistry at high
temperature/high pressure. Orig Life Evol Biosph in the ▶ interstellar medium and in comets.
22:43–65
Stribling R, Miller S (1987) Energy yields for hydrogen
cyanide and formaldehyde syntheses: the hydrogen
cyanide and amino acid concentrations in the primitive
History
ocean. Orig Life Evol Biosph 17:261–273
Walker J (1964) Formaldehyde, 3rd edn. Rheinhold, Formamide was detected for the first time in 1971
New York in the interstellar medium by Rubin et al. and in
Weber A (1998) Prebiotic amino acid thioester synthesis:
thiol-dependent amino acid synthesis from formose
1997 in the bright comet C/1995 O1 (Hale-Bopp)
substrates (formaldehyde and glycolaldehyde) and by Mehringer et al.
ammonia. Orig Life Evol Biosph 28:259–270
Weber A (2002) Chemical constraints governing the ori-
gin of metabolism: the thermodynamic landscape of
carbon group transformations under mild aqueous con- See Also
ditions. Orig Life Evol Biosph 32:333–357
Woon DE (2002) Modeling gas-grain chemistry with
quantum chemical cluster calculations. ▶ Comet
I. Heterogeneous hydrogenation of CO and H2CO on ▶ Interstellar Medium
icy grain mantles. Astrophys J 569:541–548 ▶ Molecules in Space
Zuckerman B, Buhl D, Palmer P, Snyder LE (1970) Obser- ▶ Peptide
vation of interstellar formaldehyde. Astrophys J
160:485–506
Formamide Barks HL, Buckley R, Grieves GA, Di Mauro E, Hud NV,

Orlando TM (2010) Guanine, adenine, and hypoxan-
thine production in UV-irradiated formamide solu-
Didier Despois tions: relaxation of the requirements for prebiotic
Laboratoire d’Astrophysique de Bordeaux, purine nucleobase formation. Chem Biol Chem
CNRS-Université de Bordeaux, France 11(9):1240
Mehringer D, Colom P, Benford D, Bockelee-Morvan D,
Despois D, Paubert G, Germain B, Biver N,
Synonyms Crovisier J, Gautier D, Gerard E, Rauer H, Lis DC,
Phillips TG, Moreno R, Davies JK, Dent WRF,
Methanamide; NH2CHO Owens A, Oosterbroek T, Orr A, Parmar AN,
Formation of Planetesimals: The Building Blocks of Planets 873
Antonelli LA, Fiore F, Maccarone MC, Piro L (1997) collisions with equal-sized bodies. This stage of
Comet C/1995 O1 (Hale-Bopp). IAU Circular planet formation may extend to as large as
No. 6614, 1
Rubin RH, Swenson GW Jr, Benson RC, Tigelaar HL, 1,000 km in size, depending on the strength of
Flygare WH (1971) Microwave detection of interstel- the gas turbulence which is responsible for induc-
lar formamide. Astrophys J Lett 169:L39 ing high planetesimal-planetesimal speeds.
Saladino R, Crestini C, Ciciriello F, Costanzo G, Di
Mauro E (2006) About a formamide-based origin of
informational polymers: syntheses of nucleobases and
favourable thermodynamic niches for early polymers. Overview
OLEB 36:523–531
Planet formation takes place in ▶ protoplanetary
disks of gas and dust which surround young stars.
The embedded dust and ice particles represent only F
Formamine approximately 1 % of the mass of the protoplane-
tary disk, but their structure formation through
▶ Hexamethylenetetramine collisions and sticking is responsible for the growth
of planetary bodies.
The formation of planetesimals is an impor-
tant step in the growth of planetary systems.
Formation of Planetesimals: The Planetesimals are the building blocks of the
Building Blocks of Planets rocky terrestrial planets (Mercury, Venus, Earth,
and Mars) as well as the cores of the gas giants
Anders Johansen (Jupiter and Saturn) and the ice giants (Uranus
Lund University, Lund, Sweden and Neptune). The ultimate outcome of the plan-
etesimal formation stage is the production of
planetesimals of sufficient sizes to obtain
Keywords gravitational cross sections much larger than
their physical cross sections, hence accelerating
Dust; Particle concentration; Pebbles; Planet for- the growth toward fully fledged planets.
mation; Planetesimals; Protoplanetary disks; The gravitational cross section can here refer to
Streaming instability; Turbulence either the ability to accrete other planetesimals
(Levison et al. 2010) or to accrete mm–cm-
sized pebbles facilitated by the strong gas drag
Definition felt by such small particles (Lambrechts and
Johansen 2012).
Planetesimals are traditionally defined as solid The lack of ▶ gravitational focusing in the
objects (rocky or icy or a combination of both) planetesimal formation stage means that direct
whose internal strength is dominated by self- sticking is key to particle growth. However, dust
gravity rather than material strength. This corre- particles encounter many barriers and bottlenecks
sponds to bodies of approximately 100 m to 1 km in their growth toward planetesimals, as bounc-
in size (Benz 2000). However, this definition ing and fragmentation become increasingly likely
does not take into account the role of the body with increased size (Zsom et al. 2010). Particles
in the planet-building process. Planetesimals of of sizes from mm to several m are nevertheless
km in size are vulnerable to erosion and fragmen- concentrated in the turbulent flow of the gas, and
tation in collisions with other planetesimals at the the overdensities can become high enough for the
speeds relevant in protoplanetary disks (Ida collective gravity to be dominant (Johansen
et al. 2008). One can instead define the planetes- et al. 2007). This leads to the formation of large
imal formation stage as the growth of bodies to planetesimals (100–1,000 km radii) by gravita-
sufficient sizes to be insensitive to disruption in tional collapse of particle filaments and clumps.
874 Formation of Planetesimals: The Building Blocks of Planets
Basic Methodology bouncing barrier (Zsom et al. 2010), concentrate

between the eddies (Cuzzi et al. 2001). On large
Planetesimal formation can be studied scales where Coriolis forces from the rotating
observationally, experimentally, or theoretically. disks are dominant, large-scale vortices and axi-
Important constraints on the planetesimal symmetric pressure bumps arise spontaneously in
formation process are obtained from studying the turbulent flow (Johansen et al. 2009) or near
the structure and dynamics of minor bodies in transitions in the turbulent viscosity (Lyra
the solar system – asteroids, Kuiper belt objects, et al. 2008). These geostrophic high-pressure
and comets – or from studying debris disks structures concentrate large particles, of typical
around other stars. In the latter, the observed sizes from 10 cm to 1 m.
dust is produced through collisions within an Active concentration occurs through the
(unseen) planetesimal population. In the streaming instability (Youdin and Goodman
Solar System measurements of size distributions, 2005). This hydrodynamical instability arises in
spectral classes and binarity have proven the coupled motion of particles and gas, from the
useful for developing planetesimal formation free energy present between the inward-flowing
models. particles and the outward-flowing gas in the
The outcome of collisions between particles of midplane. The gas in the protoplanetary disk is
different sizes can be measured in the laboratory. slightly pressure supported in the radial direction,
Particle aggregates are typically built from since the gas is denser and hotter close to the star.
micrometer-sized silicate monomers. These The pressure support acts like a reduced gravity
aggregates can then collide with each other and on the gas and causes the gas to orbit a bit more
the outcome is measured in terms of sticking, slowly than the Keplerian speed, by approxi-
bouncing, or fragmentation (G€uttler et al. 2010), mately 50 m/s. Solid particles do not sense the
depending on the collision speed. pressure gradient force from the gas and would
Knowing the collision outcome for a wide orbit at the Keplerian speed in absence of friction,
range of input parameters, the growth of but the headwind of the slower moving gas drains
populations of dust particles can be modeled angular momentum from the particles and causes
on a computer. The collision speed depends them to drift toward the star. The drift speed
on the sizes of the colliding particles. For peaks for typical particle sizes of 10 cm to 1 m,
small particles, the Brownian motion in the inner few AU of the protoplanetary disk,
dominates, while larger particles have with a drift timescale as low as a few hundred
increasing contributions from the turbulent years. Regions of slightly increased particle den-
gas, peaking at approximately 5–50 m/s for sity drift more slowly toward the star as the gas
m-sized particles under typical turbulent there is accelerated toward the Keplerian speed
conditions. by the drag force from the particles. The stream-
Hydrodynamical simulations are used to ing instability causes an exponential increase of
investigate the concentration of particles in the the density of the filament as isolated, fast-
turbulent gas flow of the ▶ protoplanetary disk. drifting particles continue to pile up in the
Particle concentration mechanisms can be overdense region.
divided into two categories. Passive concentra-
tion occurs as particles pile up in regions of high
gas pressure. On small scales of the turbulent Key Research Findings
flow, these regions reside between swiftly rotat-
ing turbulent eddies (which are themselves low Dust growth models have identified a number of
pressures). Small particles, under some condi- bottlenecks in the planetesimal formation pro-
tions similar in sizes to those stuck at the cess. The electrostatic barrier describes the
Formation of Planetesimals: The Building Blocks of Planets 875
inability for dust grains to grow beyond a few magnetorotational instability (Balbus and
10 mm, as negatively charged aggregates repel Hawley 1991). Computer simulations of
each other (Okuzumi 2009). The bouncing bar- magnetorotational turbulence in accretion disks
rier sets in at sizes from 0.1 to 1 mm, as show the emergence of large-scale pressure
compactified dust aggregates of these sizes will bumps which trap dm–m-sized dust particles
bounce rather than stick if collision speeds are (Johansen et al. 2009). The streaming instability
higher than a few mm per second (Zsom was discovered by linear stability analysis by
et al. 2009). The fragmentation barrier sets in at Youdin and Goodman (2005). The process of
collision speeds higher than 1 m/s, although piling up isolated particles into overdense fila-
growth can still be obtained if small impactors ments leads to very high particle densities, as
hit much larger targets at speeds up to 25 m/s high as 10,000 times the local gas density
(Wurm et al. 2005). Finally the formidable radial (Johansen et al. 2012). F
drift barrier requires that growth must be faster The growth of particles by sticking and col-
than the radial drift of particles (Weidenschilling lapse of overdense particle filaments into plane-
1977). Even if the radial drift barrier could be tesimals are not mutually exclusive. Particles
somehow overcome, the fragmentation barrier must grow to sizes of at least mm in order to be
has second branch for km-scale planetesimals able to concentrate in the turbulent gas, and some
whose collision speed can be excited to super- mechanisms like vortices and pressure bumps
escape values by the turbulent gas (Ida seem to require particles as large as 10 cm to
et al. 2008). begin with. Particle growth also does not stop at
While ideas exist for circumventing the many the collapse phase. Particle densities reach very
growth barriers, e.g., by invoking sticky organics high values during the contraction to solid densi-
(Kouchi et al. 2002) or dust rims to enhance ties and hence collision rates are high. The out-
sticking between the mm-sized chondrules comes of these collisions determine the sizes of
which are found in meteorites (Ormel the planetesimals which emerge from the
et al. 2008), any growth must compete with the contracting clouds. Nesvorny et al. (2010) inves-
radial drift of the particles. Global models of tigated the outcome of collapsing pebble clouds
particle coagulation have only managed to pro- and found that an initial mean rotation of the
duce planetesimals by sticking either in the inner- cloud (which could be inherited from the overall
most part of the disk (within 1 AU) or by rotation of the disk) leads typically to the forma-
assuming high sticking threshold speeds and tion of a binary planetesimal surrounded by par-
invoking pressure bumps to stop the radial drift ticles clumps which can be interpreted as a swarm
(Brauer et al. 2008). of smaller planetesimals. This model agrees with
Another path to planetesimal formation is the observation that the classical cold component
through the concentration of particles in the tur- of the Kuiper belt – the most pristine planetesi-
bulent gas flow, followed by a brief phase of mals in the Solar System which have undergone
gravitational contraction and collapse. Large- very little collisional and dynamical
scale axisymmetric pressure bumps were identi- evolution – have a very high binary fraction.
fied already by Whipple (1972) as traps of
drifting particles. Barge and Sommeria (1995)
explored dust trapping in anticyclonic Applications
vortices – structures which could arise in proto-
planetary disks around at transitions from weak to Results from planetesimal formation models
strong turbulence (Lyra et al. 2008) or at the edge can be used as input for understanding the
of gaps produced by giant planets. Turbulence in formation of planetary systems. The birth sizes
protoplanetary disks may be driven by the of planetesimals can be applied to the
876 Formation of Planetesimals: The Building Blocks of Planets
asteroid belt to understand the origin of the ▶ Small Solar System Body
current size distribution of asteroids as well as ▶ Solar Nebula
the emergence of the major asteroid ▶ Turbulence (Planetary Disks)
classes. Water delivery to Earth is thought to ▶ Vortex, Vortices
have happened via collisions with icy
planetesimals, so the origin of life on Earth is
deeply connected to the formation of
planetesimals. Balbus SA, Hawley JF (1991) A powerful local shear
instability in weakly magnetized disks. I – linear anal-
ysis. II – nonlinear evolution. Astrophys
Future Directions J 376:214–233
Barge P, Sommeria J (1995) Did planet formation begin
inside persistent gaseous vortices? Astron Astrophys
An important priority for planetesimal formation 295:L1–L4
studies in the future is to develop models of the Benz W (2000) Low velocity collisions and the growth of
growth of dust particles by sticking which take planetesimals. Space Sci Rev 92:279–294
Brauer F, Henning T, Dullemond CP (2008) Planetesimal
into account the concentration and gravitational formation near the snow line in MRI-driven turbulent
collapse into planetesimals of a wide range of protoplanetary disks. Astron Astrophys 487:L1–L4
sizes. While many aspects of particle growth G€uttler C, Blum J, Zsom A, Ormel CW, Dullemond CP
and concentration are well understood, the single (2010) The outcome of protoplanetary dust growth:
pebbles, boulders, or planetesimals? I. Mapping the
most important parameter in determining which zoo of laboratory collision experiments. Astron
processes are dominant is the strength of the Astrophys 513:A56
turbulence in the gas. Strong background turbu- Ida S, Guillot T, Morbidelli A (2008) Accretion and
lence suppresses the streaming instability so that destruction of planetesimals in turbulent disks.
particles must grow to sizes of at least 10 cm in Johansen A, Oishi JS, Mac Low M-M, Klahr H,
order to concentrate in large-scale pressure Henning T, Youdin A (2007) Rapid planetesimal for-
bumps and vortices. The streaming instability mation in turbulent circumstellar disks. Nature
can concentrate particles down to cm sizes if the 448:1022–1025
Johansen A, Youdin A, Klahr H (2009) Zonal flows and
turbulence is moderate or weak. Thus, an long-lived axisymmetric pressure bumps in magnetor-
increased comprehension of the presence and otational turbulence. Astrophys J 697:1269–1289
strength of turbulence in protoplanetary disks Johansen A, Youdin AN, Lithwick Y (2012) Adding par-
will be crucial for understanding how planetesi- ticle collisions to the formation of asteroids and Kuiper
belt objects via streaming instabilities. Astron
mals form in protoplanetary disks. Astrophys 537:A125
Kouchi A et al (2002) Rapid growth of asteroids owing to
very sticky interstellar organic grains. Astrophys
J 566:L121–L124
See Also Lambrechts M, Johansen A (2012) Rapid growth of
gas-giant cores by pebble accretion. Astron Astrophys
▶ Asteroid 544:A32
Levison HF, Thommes E, Duncan MJ (2010) Modeling
▶ Coagulation in Planetary Disks
the formation of giant planet cores. I. Evaluating key
▶ Core Accretion, Model for Giant Planet processes. Astron J 139:1297–1314
Formation Lyra W, Johansen A, Klahr H, Piskunov N (2008) Embryos
▶ Gas drag grown in the dead zone. Assembling the first protoplan-
▶ Kuiper Belt etary cores in low mass self-gravitating circumstellar
disks of gas and solids. Astron Astrophys 491:L41–L44
▶ Meter-Size Catastrophe Nesvorny D, Youdin AN, Richardson DC (2010) Forma-
▶ Planet tion of Kuiper belt binaries by gravitational collapse.
▶ Planet Formation Astron J 140:785–793
▶ Protoplanetary Disk Okuzumi S (2009) Electric charging of dust aggregates
and its effect on dust coagulation in protoplanetary
▶ Runaway Growth disks. Astron J 698:1122–1135
Formose Reaction 877
Ormel CW, Cuzzi JN, Tielens AGGM (2008) Co-accretion formamide. Melting point, 8.4 C; boiling
of chondrules and dust in the solar nebula. Astron point, 101 C; density, 1.22 g cm3.
J 679:1588–1610
Weidenschilling SJ (1977) Aerodynamics of solid bodies
in the solar nebula. Mon Not R Astron Soc 180:57–70
Wurm G, Paraskov G, Krauss O (2005) Growth of plane- References and Further Reading
tesimals by impacts at 25 m/s. Icarus 178:253–263
Youdin AN, Goodman J (2005) Streaming instabilities in Garrison WM, Morrison DC, Hamilton JG, Benson AA,
protoplanetary disks. Astrophys J 620:459–469 Calvin M (1951) Reduction of carbon dioxide in aque-
Zsom A, Ormel CW, G€ uttler C, Blum J, Dullemond CP ous solutions by ionizing radiation. Science
(2010) The outcome of protoplanetary dust growth: 114:416–418
pebbles, boulders, or planetesimals? II. Introducing
the bouncing barrier. Astron Astrophys 513:A57
F
Formic Acid Methyl Ester
Formic Acid
▶ Methyl Formate
Kensei Kobayashi
Hodogaya-ku, Yokohama, Japan
Formonitrile
Synonyms ▶ Hydrogen Cyanide
Methanoic acid
Formose Reaction
Definition
Formic acid is a carboxylic acid, whose chemical Earth–Life Science Institute (ELSI), Tokyo
formula is HCOOH (Fig. 1). It has a dissociation Institute of Technology, Meguro–ku, Tokyo,
constant in water (pKa) of 3.74. It dissolves read-
Japan
ily in water and is slightly soluble in hydrocar- Institute for Advanced Study, Princeton, NJ,
bons. It is mostly present as dimers in USA
hydrocarbon solvents and in the gas phase. It
can react as a reducing agent. It dissociates to Washington, DC, USA
carbon monoxide and water upon heating or by Center for Chemical Evolution, Georgia Institute
the addition of sulfuric acid. In an early report on
the abiotic synthesis of organic compounds, Gar-
rison et al. (1951) reported that formic acid was
produced by helium ion irradiation of an aqueous
Keywords
solution containing CO2 and Fe2+. Formate is
also produced by the hydrolysis of HCN or Autocatalysis; Carbohydrate; Formaldehyde;
Formose; Ribose; RNA World
Formic Acid, O
Fig. 1 Formic acid
C Synonyms
H OH
Butlerow reaction
878 Formose Reaction
Definition Reaction Mechanism

The mechanism of the reaction has been studied
The formose reaction, discovered by Butlerow in intensively, and the complexity of the product
1861, is a complex autocatalytic set of mixture is notorious (Breslow 1959). The reac-
condensation reactions of formaldehyde to tion is typically carried out from concentrated
yield sugars and other small sugar-like mole- HCHO under alkaline conditions in the presence
cules. The reaction is particularly noteworthy in of divalent metal ions, such as calcium, magne-
the context of astrobiology and prebiotic chemis- sium, or lead, and is preceded by an induction
try in that it could serve as a potential abiotic period which can be shortened by the addition of
source of carbohydrates, in particular ribose, glycolaldehyde or other aldehydes as initiators.
which could be important for the origin of an Reaction does not proceed at an appreciable rate
RNA World. under neutral conditions at room temperature. At
high pH, it is sluggish in the absence of catalysts
such as divalent metal ions (Orgel 2000). It has
Overview been difficult to decipher whether the reaction
requires divalent cations or higher aldehydes to
The formose reaction is an autocatalytic reaction initiate. It has been reported that carbohydrates
discovered by Butlerow (1861). It involves the and other trace impurities in commercial formal-
formation of sugars, polyols and hydroxy acids dehyde solutions or paraformaldehyde present in
from ▶ formaldehyde in a series of carbon-to- ppm quantities may be the cause of the autoca-
carbon condensations, as opposed to carbon-to- talysis observed in some of the formose reactions
oxygen condensations of HCHO to form studied to date (Socha et al. 1980). For example,
▶ polyoxymethylene. Formose is a contraction paraformaldehyde sublimed into Ca(OH)2 sus-
of formaldehyde and the suffix -ose, denoting a pension was not transformed to sugars via the
sugar. In fact, many biological sugars have formose reaction, but only to methanol and for-
empirical formulas of the form (CH2O)n, for mate by the base-catalyzed Cannizzaro reaction,
example, glucose, (CH2O)6, and ▶ ribose, and a 3 ppm trace of glycolaldehyde was suffi-
(CH2O)5. The formose reaction may be a mech- cient to initiate conventional autocatalysis
anism for the prebiotic synthesis of sugars from (Socha et al. 1980). However, at neutral or
formaldehyde, of relevance to the origin of an slightly basic conditions, redistilled formalde-
▶ RNA World (Gesteland and Atkins 1993), hyde solutions undergo the classical reaction
early nucleic acids based on alternative sugars (at 80 ) in the presence of several minerals,
in a potential “pre-RNA World” (Joyce et al. while no reaction is observed in their absence
1987), as well as other sugar-based models for (Schwartz and de Graaf 1993a; see below).
the origin of life (Weber 1997, 2001). The The reaction involves a series of intermediate
formose reaction is one of the few chemical sys- steps including aldol reactions, reverse aldol
tems proposed to be close to an autocatalytic reactions, and isomerization reactions.
proto-metabolic cycle (Orgel 2000). A number of intermediates have been identified
Both formaldehyde and ▶ glycolaldehyde including glycolaldehyde, glyceraldehyde, dihy-
have been observed spectroscopically in outer droxyacetone, and tetroses. Figure 1 shows a
space in both the ▶ interstellar medium and in simplified proposed mechanism for the reaction
comets (Hollis et al. 2000). Formaldehyde and (Breslow 1959).
sugar-related compounds, such as ▶ glycerol, The reaction is proposed to begin by the appar-
which could be derived from this reaction are ently kinetically slow reaction of two formalde-
abundant in some carbonaceous chondrites hyde molecules to make glycolaldehyde.
(Cooper et al. 2001); thus, the formose reaction Glycolaldehyde then reacts via an aldol conden-
could be a cosmically common abiotic mecha- sation with another formaldehyde molecule to
nism for sugar synthesis. yield DL-glyceraldehyde. Isomerization of
Formose Reaction, Fig. 1 A simplified scheme for the not specified, but in general all isomers are formed. Side
formose reaction. The stereochemistry of the asymmetric reactions leading to branched-chain molecules complicate
carbon atoms (marked with an asterisk in the diagram) is the cycle and divert molecules from it
glyceraldehyde forms dihydroxyacetone (DHA), minerals, though the ones that do exist suggest
which can react with glycolaldehyde to form a 2- that HCHO-clay adsorption equilibria are not
ketopentose. This 2-ketopentose may then isom- especially high, which is also true for low molec-
erize to yield various isomeric pentoses. DHA ular weight POMs (Parfitt and Greenland 1970).
can also react with formaldehyde to produce a Adsorption equilibria may be higher for clays
2-ketotetrose which can isomerize to give an such as illite and kaolinite (Chandra and De
aldotetrose. At this step, the aldotetrose can 1983).
react via a retro-aldol reaction to give two mole- Many minerals appear to catalyze the formose
cules of glycolaldehyde. Due to the generation of reaction, for example, various clays, apatite, and
two molecules of glycolaldehyde from the input calcite were found to be catalysts (Gabel and
of one, the reaction becomes autocatalytic. Ponnamperuma 1967; Reid and Orgel 1967;
Cairns-Smith et al. 1972; Schwartz and de Graaf
Mineral Interactions 1993b). The reaction can be carried out at lower
Mineral adsorption may have been an important pH and in the absence of glycolaldehyde initiator
sink or concentration mechanism for HCHO and in the presence of some minerals (Schwartz and
sugars. Mineral surfaces and other inorganic spe- de Graaf 1993a). Certain mineral types, in partic-
cies can also have significant effects on the ular ▶ borates and silicates, stabilize intermedi-
course of the reaction, as both general and selec- ates in the reaction pathway at high pH and might
tive catalysts for synthesis and degradation for significantly alter the product mixture under these
the reaction products. There have been relatively conditions (Ricardo et al. 2004; Lambert
few studies of the adsorption of HCHO to et al. 2010). This may be particularly significant
given the wide distribution of silicates in the solar act as stabilizers (Ricardo et al. 2004; Lambert
system. Double-layer hydroxide minerals have et al. 2010).
been shown to be catalysts for aldol condensa-
tions under very dilute conditions (Arrhenius Photocatalysis and Reaction with CH3CHO
et al. 1994). The photochemical synthesis of formaldehyde
Mineral effects have generally been studied from ultraviolet irradiation of CO2 and water is
using high concentrations of HCHO (0.01 M or quite robust (Pinto et al. 1980). Although its
higher) or other aldehydes; it remains unknown hydrate has a low UV cross section, HCHO may
whether minerals affect the reaction of more have been prone to photoreaction in primitive
dilute HCHO (see below). surface waters. Schwartz and De Graaf found
that UV irradiation of dilute solutions of formal-
Competing Chemistry dehyde produced acetaldehyde (CH3CHO) in sig-
nificant yield, as well as very high yields of
Cannizzaro Reactions pentaerythritol (Fig. 2) (Schwartz and de Graaf
At high concentrations, HCHO undergoes a dis- 1993a). Other nonsugar products have been
proportionation reaction to form formate and observed from photochemical formose reactions
methanol (Eq. 1) (Walker 1964), which reduces by others (Shigemasa et al. 1977).
the amount of HCHO available to form sugars. Significant quantities of acrolein
These reactions are acid, base, and metal cata- (CH2=CHCHO) have also been detected in
lyzed (Walker 1964). The reaction reportedly spark discharge experiments (van Trump and
shows third-order kinetics. It is not clear whether Miller 1972). The DG for the condensation of
it would occur significantly in dilute solution. CH3CHO and HCHO in the gas phase was esti-
mated at 4.44 kcal mol1 and the Keq as 1640
2HCHO þ H2 O ! CH3 OH þ HCOOH (1) (Malinowski et al. 1963). Thus, the overall reac-
tion is thermodynamically favorable.
Oligomerization to Polyoxymethylene Acrolein is an intermediate in the industrial
A series of low molecular weight polymers, synthesis of pentaerythritol from concentrated
polyoxymethylenes (POMs) (of formula HO HCHO and CH3CHO in aqueous alkaline solu-
(CH2O)nH) as well as small six- and eight- tion (Berlow et al. 1958). HCHO reacts with
membered cyclic oligomers, form readily in neu- CH3CHO at much lower concentrations, to give
tral concentrated aqueous HCHO solutions. This acrolein among other products, than the formose
polymerization is highly concentration depen- reaction has been observed to occur (Cleaves
dant. A 5 wt% ( 1.3 M) aqueous HCHO solution 2003). The synthesis of acrolein from equimolar
contains only 14 % POM dimer, 3 % trimer, and HCHO and CH3CHO was found to be fairly inde-
0.5 % tetramer. Polymerization equilibria would pendent of pH between pH 7 and 11 and rela-
be much lower for the prebiotic oceanic concen- tively independent of concentration between of
trations of HCHO which have been estimated. 104 and 1 M. In contrast, glycolaldehyde and
glyceraldehyde syntheses were found to be
Sugar Degradation extremely concentration dependent. At 103 M
Under the basic reaction conditions which allow concentration of equimolar HCHO and CH3CHO
the formose reaction to occur, sugar decomposi- and below, neither glycolaldehyde nor glyceral-
tion into tars (Reid and Orgel 1967; Shapiro dehyde was detected. The same is presumably
1988), furans, and other low molecular weight true of the higher sugars.
degradation products (De Bruijn et al. 1986; Coo- It has been demonstrated that acrolein is one of
per et al. 2001) is also facilitated (Larralde the few compounds with which the nucleobases
et al. 1995). This has been shown to be less of RNA/DNA will readily react (Nelsestuen
significant depending on the presence of high 1980). Thus, acrolein could have been an impor-
concentrations of inorganic species which may tant sink for the nucleobases in the prebiotic
Formose Reaction, Fig. 2 Scheme for the photochemical synthesis of CH3CHO, acrolein, and pentaerythritol from
UV irradiation of HCHO
environment and a significant hindrance to the If the concentration and reaction of HCHO on
start of an RNA World. the primitive Earth were difficult because of all of
the potential competing geochemical sinks,
Environmental Limitations sugars may have been derived from meteoritic
The efficiency of the formose reaction in a prebi- input. Polyols such as glycerol and ribitol have
otic environment is highly dependent on the rate been identified in the Murchison meteorite,
of HCHO production and delivery to the primi- although the only actual sugar identified was
tive oceans (Pinto et al. 1980). The reaction has DHA (Cooper et al. 2001). This can potentially
been observed to occur with HCHO as dilute as be explained by the instability of sugars over the
0.01 M (Reid and Orgel 1967; Schwartz and De 4 billion years since the Murchison parent body
Graaf 1993a), although there are accounts of its formed and the observed organic synthesis pre-
reaction as dilute as 103 M (Gabel and sumably occurred. The initial synthesis to form
Ponnamperuma 1967). These concentrations sugars may have occurred relatively rapidly on
may have been difficult to achieve in the bulk the parent body, while only those compounds
early oceans. As HCHO became concentrated, stable enough to survive until the present day
side reactions may have established a low are still detectable. The detection of these com-
steady-state concentration regardless of the pro- pounds does however present an interesting par-
duction rate. Given the observed photochemical adox for prebiotic chemistry. If the polyols and
synthesis of pentaerythritol described above, it their precursors were generated by the same
may be unlikely that bulk oceanic concentrations aqueous phase chemistry which produced the
of HCHO could have been much higher than amino and hydroxy acids, as well as the purines
103 M. and pyrimidines which have been detected in the
Murchison meteorite (Pizzarello 2004), appar- temperatures (Larralde et al. 1995). The eutectic
ently the suggested inhibition of HCHO chemis- freezing and thawing of precursor ice-grain
try by HCN and vice versa (Schlesinger and organics such as HCHO in carbonaceous chon-
Miller 1973) is not a genuine problem. The ratio drites may have allowed for the synthesis of
of glycolic acid to glycine in Murchison suggests sugars on these bodies (Cooper et al. 2001). The
that NH3 concentrations were fairly high in the frozen clay model proposed by Lahav and Chang
parent body (Peltzer et al. 1984). These quantities (1976) may warrant careful reconsideration.
also suggest that the reactions which form HMT
and glycolonitrile are not limiting to either purine Concentration by Evaporation
or sugar synthesis and that slow kinetic effects Evaporative concentration is a plausible geo-
may be more important than the initial rapid chemical mechanism for concentrating nonvola-
equilibrium obtained. tile species. The equilibrium constant for the
The prebiotic oceanic concentrations of most dimerization of HCHO to give glycolaldehyde
precursor organic compounds such as HCHO and is not known, but is of considerable interest,
NH3 were also likely quite low (Stribling and since while HCHO is volatile, glycolaldehyde is
Miller 1987). It is necessary to consider the geo- not and could thus be concentrated by evapora-
chemical processes which might have concen- tion. A rough calculation using the free energy of
trated HCHO sufficiently to allow the formose the aldol reaction of HCHO given by Weber
reaction to occur. It is worth considering what (2002) suggests the equilibrium constant is
happens to dilute solutions of more complex mix- 40, but this does not take into account various
tures of HCN, HCHO, and other aldehydes, NH3, kinetic factors which may limit the reaction. This
urea, nitrate, nitrite, sulfides, ferrous iron, salts, simple reaction is worthy of further investigation
etc., as they are evaporated in the presence of UV from the standpoint of prebiotic chemistry, espe-
and visible light over mineral surfaces, or as they cially given the presence of glycolaldehyde in
are frozen from dilute solution under visible interstellar space (Hollis et al. 2000).
and/or UV irradiation.
Hydrothermal Vents
Possible Geological Settings for the Formose Although some hydrothermal vent environments
Reaction are characterized by rather alkaline conditions
Sugars are easily made by the reaction of HCHO which would be ideal for formose chemistry,
under certain conditions (fairly high concentra- most modern vent systems are generally charac-
tions and slightly basic to basic pH). Such condi- terized by low organic concentrations and mod-
tions of pH and concentration are likely only erate to very high temperatures and thus may not
attainable in certain geological environments be optimal sites for formose chemistry. In addi-
such as in eutectics and in evaporative environ- tion to the thermal instability of sugars, HCHO
ments after HCHO is rendered nonvolatile by does not appear to be a stable one-carbon species
reaction with other aqueous species, for example, under hydrothermal conditions (Osada
NH3, SO 3 , or H2S. et al. 2004; Seewald et al. 2006).
Given the uncertainties regarding primitive
Eutectic Freezing Earth conditions, the existence of specialized
Eutectic freezing can effectively concentrate conditions which would allow the formose reac-
dilute prebiotic reactants to form purines, pyrim- tion to occur cannot be ruled out. The synthesis of
idines, and ▶ amino acids (Sanchez et al. 1966; sugars in such environments from low initial
Levy et al. 2000; Miyakawa et al. 2002). Whether HCHO concentrations under mild conditions
HCHO can also be concentrated via eutectic might be extremely facile and warrants further
freezing to form sugars is unknown, but plausi- investigation, as does formose synthesis in the
ble. One significant advantage of this process is presence of congeners such as HCN, NH3, and
that sugars are generally much more stable at low inorganic compounds such as borate, sulfide, and
sulfite and the impact of UV and visible radiation Gabel N, Ponnamperuma C (1967) Model for origin of
on this chemistry. monosaccharides. Nature 216:453–455
Gesteland R, Atkins J (1983) The RNA world: the nature
As mentioned above, the perceived impor- of modern RNA suggests a prebiotic RNA world
tance of the formose reaction rests mainly on its (Monograph/Cold Spring Harbor Laboratory, No 24)
role in sugar synthesis which is important in the Gesteland RF, Atkins JF (1993) The RNA world: the nature
context of abiotic nucleic acid synthesis. Sugar of modern RNA suggests a prebiotic RNA world. Cold
Spring Harbor Laboratory Press, Cold Spring Harbor
synthesis, of course, is merely one step on the way Hollis J, Lovas F, Jewell P (2000) Interstellar
to nucleoside, nucleotide, and nucleic acid syn- glycolaldehyde: the first sugar. Astrophys J 540:
thesis, which also may require certain specialized L107–L110
environmental conditions (Fuller et al. 1972), Joyce G, Schwartz A, Miller S, Orgel L (1987) The case
for an ancestral genetic system involving simple ana-
although there are proposed mechanisms for logues of the nucleotides. Proc Natl Acad Sci U S A
nucleotide formation which do not necessarily 84:4398–4402 F
depend on direct condensation of the nucleic Lahav N, Chang S (1976) The possible role of solid
acid base with a sugar (Powner et al. 2009). surface area in condensation reactions during chemical
evolution: reevaluation. J Mol Evol 8:357–380
Lambert JB, Gurusamy-Thangavelu SA, Ma K (2010) The
Silicate-Mediated formose reaction: bottom-up syn-
See Also thesis of sugar silicates. Science 327:984–986
Larralde R, Robertson M, Miller S (1995) Rates of decom-
position of ribose and other sugars: implications for
▶ Carbohydrate chemical evolution. Proc Natl Acad Sci U S A
▶ Formaldehyde 92:8158–8160
▶ Ribose Levy M, Miller S, Brinton K, Bada J (2000) Prebiotic
▶ RNA World synthesis of adenine and amino acids under Europa-
like conditions. Icarus 145:609–613
Malinowski S, Basinski S, Szczepanska (1963) Ann Soc
Chim Polonorum 37:977–982
References and Further Reading Miyakawa S, Cleaves H, Miller S (2002) The cold origin
of life: B. Implications based on pyrimidines and
Arrhenius T, Arrhenius G et al (1994) Archean geochem- purines produced from frozen ammonium cyanide
istry of formaldehyde and cyanide and the oligomeri- solutions. Orig Life Evol Biosph 32:209–218
zation of cyanohydrin. Orig Life Evol Biosph Nelsestuen GL (1980) Origin of life: consideration of
24(1):1–17 alternatives to proteins and nucleic acids. J Mol
Berlow E, Barth RH, Snow JE (1958) The pentaery- Evol 15(1):59–72
thritols. Reinhold Publishing, NY Orgel LE (2000) Self-organizing biochemical cycles.
Breslow R (1959) On the mechanism of the formose PNAS 97(23):12503–12507
reaction. Tetrahedron Lett 21:22–26 Osada M, Watanabe M, Sue K, Adschiri T, Arai K (2004)
Butlerow A (1861) Formation synthétique d’une sub- Water density dependence of formaldehyde reaction in
stance sucrée. Comp Rend Acad Sci 53:145–147 supercritical water. J Supercrit Fluids 28:219–224
Cairns-Smith A, Ingram P, Walker G (1972) Formose Parfitt R, Greenland D (1970) The adsorption of poly
production by minerals: possible relevance to the ori- (ethylene glycols) on clay minerals. Clay Miner
gin of life. J Theor Biol 35:601–604 8:305–315
Chandra K, De S (1983) Adsorption of formaldehyde by Peltzer E, Bada J, Schlesinger G, Miller S (1984) The
clay minerals in presence of urea and ammonium sul- chemical conditions on the parent body of the Murchi-
fate in aqueous system. Indian J Agr Chem son meteorite: some conclusions based on amino,
16:239–245 hydroxy and dicarboxylic acids. Adv Space Res
Cleaves H (2003) The prebiotic synthesis of acrolein. 4:69–74
Monatsh Chem 134:585–593 Pinto J, Gladstone G, Yung Y (1980) Photochemical pro-
Cooper G, Kimmich N, Belisle W, Sarinana J, Brabham K, duction of formaldehyde in Earth’s primitive atmo-
Garrel L (2001) Carbonaceous meteorites as a source sphere. Science 210:183–185
of sugar-related organic compounds for the early Pizzarello S (2004) Chemical evolution and meteorites: an
Earth. Nature 414:879–883 update. Orig Life Evol Biosph 34:25–34
De Bruijn J, Kieboom A, Van Bekkum H (1986) Reactions Powner MW, Gerland B, Sutherland JD (2009) Synthesis
of monosaccharides in aqueous alkaline solutions. of activated pyrimidines ribonucleotides in
Sugar Tech Rev 13:21–52 prebiotically plausible conditions. Nature
Fuller W, Sanchez R, Orgel L (1972) Studies in prebiotic 459:239–242
synthesis VII. J Mol Evol 1:249–257
884 Formyl Cation
Reid C, Orgel L (1967) Synthesis of sugars in potentially Definition

prebiotic conditions. Nature 216:455
Ricardo A, Carrigan M, Olcott A, Benner S (2004) Borate
minerals stabilize ribose. Science 303:196 The triatomic ion HCO+ is one of the most abun-
Sanchez R, Ferris J, Orgel L (1966) Conditions for purine dant molecular ions in dense interstellar clouds. It
synthesis: did prebiotic synthesis occur at low temper- is a key intermediary in the production of ▶ car-
atures? Science 153:72–73 bon monoxide, CO, which is in turn the most
Schlesinger G, Miller S (1973) Equilibrium and kinetics of
glyconitrile formation in aqueous solution. J Am Chem abundant constituent of these regions after
Soc 95:3729–3735 molecular hydrogen (H2). The abundance ratio
Schwartz A (1983) Chemical evolution: the first stages. of HCO+ to its deuterated counterpart, DCO+,
Naturwissenschaften 70:373–377 can provide a measure of the electron density in
Schwartz A, De Graaf R (1993a) The prebiotic synthesis of
carbohydrates: a reassessment. J Mol Evol 36:101–106 molecular clouds. HCO+ is a linear, closed-shell
Schwartz AW, de Graaf RM (1993b) Tetrahedron Lett molecule, so that its pure rotational spectrum
34:2201 consists of a series of harmonically related lines,
Seewald JS, Zolotov M, McCollom T (2006) Experimen- with the lowest transition in the 3 mm
tal investigation of single carbon compounds under
hydrothermal conditions. Geochim Cosmochim Acta wavelength band.
70:446–460
Shapiro R (1988) Prebiotic ribose synthesis: a critical
analysis. Orig Life Evol Biosph 18:71–85 History
Shigemasa Y, Matsuda Y, Sakazawa C, Matsuura
T (1977) Formose reactions II. The photochemical
formose reaction. Bull Chem Soc Jpn 50:222–226 In 1970, L. Snyder and D. Buhl reported the
Socha RF, Weiss AH, Sakharov MM (1980) Autocatalysis detection of an unidentified emission line in the
in the formose reaction. React Kinet Catal Lett spectra of several interstellar ▶ molecular clouds
14(2):119–128
Stribling R, Miller S (1987) Energy yields for hydrogen and referred to the unknown carrier as “X-ogen.”
cyanide and formaldehyde syntheses: the hydrogen Laboratory measurements subsequently
cyanide and amino acid concentrations in the primitive confirmed that X-ogen was HCO+, in agreement
ocean. Orig Life Evol Biosph 17:261–273 with the theoretical predictions of
Van Trump JE, Miller SL (1972) Prebiotic synthesis of
methionine. Science 178(63):859–860 ion-molecule chemistry by W. Klemperer and
Walker J (1964) Formaldehyde, 3rd edn. Rheinhold, E. Herbst that this ion should be abundant in
New York such clouds.
Weber A (1997) Energy from redox disproportionation of
sugar carbon drives biotic and abiotic synthesis. J Mol
Evol 44:354–360
Weber A (2001) The sugar model: catalysis by amines
See Also
and amino acid products. Orig Life Evol Biosph
31:71–86 ▶ Carbon Monoxide
Weber A (2002) Chemical constraints governing the ori- ▶ Deuterium
gin of metabolism: the thermodynamic landscape of
carbon group transformations under mild aqueous con-
▶ Molecular Cloud
ditions. Orig Life Evol Biosph 32:333–357 ▶ Molecules in Space

Formyl Cation
Herbst E, Klemperer W (1974) Is X-ogen HCO+.
William M. Irvine Astrophys J 188:255–256
Synonyms Förster Resonance Energy Transfer
HCO+ ▶ FRET
Fossil 885
Forsterite Fossa, Fossae
▶ Olivine Ernst Hauber

(DLR) e.V., Institut f€ur Planetenforschung,
Berlin, Germany
Fortescue Group
Definition
Martin J. Van Kranendonk
School of Biological, Earth and Environmental Fossae are long, narrow depressions, often occur- F
Sciences, University of New South Wales, ring in arrays (definition by the International
Australia Astronomical Union; http://planetarynames.wr.
usgs.gov/jsp/append5.jsp). This is a descriptor
term for naming surface features on the ▶ terres-
trial planets, the Moon, and on icy satellites.
Definition
The 2.77–2.63 Ga Fortescue Group is a well-

preserved, thick (0.5–6 km) succession of conti-
See Also
nental flood ▶ basalts and interbedded
▶ Moon, The
sedimentary rocks deposited unconformably on
basement granite-greenstone crust of the
▶ Pilbara Craton, Western Australia. Together
with the contemporaneous Ventersdorp
basalts in South Africa, the volcanic rocks are
the oldest known continental flood basalts
(trapps). The 2.63–2.45 Ga Hamersley Group, Fossil
containing thick deposits of ▶ banded iron for-
mation, conformably overlies the group. Sedi- Emmanuelle J. Javaux
mentary rocks of the Fortescue Group contain Palaeobiogeology-Palaeobotany-
several horizons of stromatolitic carbonates, Palaeopalynology, Geology Department,
deposited in freshwater lacustrine environments. Université de Liège, Liège, Belgium
Controversial filaments showing septa have also
been observed and related to septate microbial
microfossils. Definition
A fossil is the preserved remnant of an organism

after its death. A fossil may be morphological or
See Also chemical. A fossil can have various sizes
(microscopic to macroscopic) and composition
▶ Archean Traces of Life (organic or mineral) and may represent a whole
▶ Banded Iron Formation organism, part of an organism, colonial organ-
▶ Basalt isms, or the morphological or chemical imprint of
▶ Pilbara Craton an organism or of its activity. The processes of
▶ Stromatolites fossilization are complex, and they may lead to
▶ Trapps the preservation of organisms in their original
886 Fossilization Processes
composition or partially or be completely or Archaea or of viruses has long been debated

replaced by another (organic or mineral) material due in particular to the difficulty to recognize
or preserved as a mold or cast or as traces of unambiguously such fossils in old rocks. ▶ Bio-
activity such as footprints, trails, or burrows mineralization controlled or induced by microor-
(“ichnofossil”). ganisms themselves seems to be a major process
allowing preservation of cell structures and/or of
organic molecules.
See Also
▶ Biomarkers, Morphological Overview

▶ Dubiofossil The three domains of life can, in principle, be
▶ Fossilization, Process of preserved as ▶ microfossils, depending on the
▶ Microfossils conditions of preservation and their original com-
▶ Molecular Fossils position. Most fossils in the Phanerozoic are
▶ Pseudofossil made of hard mineral parts or imprints of hard
mineral parts of metazoans or eukaryote unicel-
lular organisms (e.g., diatoms, foraminifers).
However, we focus here on processes of fossili-
zation of soft tissues and in particular of micro-
Fossilization Processes
organisms which are more relevant to
astrobiology. Interestingly, it has been widely
▶ Biosignatures, Effect of Metamorphism
suggested that microorganisms were often
involved in the fossilization of metazoan soft
tissues (e.g., Raff et al. 2006).
There are undisputed fossils of prokaryotes in
Fossilization, Process of the geological record, and some have been
assigned taxonomic names such as Girvanella
Karim Benzerara (Riding 2002). The abundance of these microfos-
Institut de Minéralogie et de Physique des sils has varied through geological time. The
Milieux Condensés, UMR 7590, CNRS, absence of microfossils has sometimes been
Université Pierre et Marie Curie & Institut de interpreted as evidence for abiotic conditions
Physique du Globe de Paris, Paris, France prevailing during the formation of a rock. For
example, the absence of microfossils in Archean
stromatolites has been used to question their
Keywords biogenicity (e.g., Grotzinger and Rothman
1996). Alternatively, the varying abundance of
Biomineralization; Diagenesis; Taphonomy microfossils has often been related to variations
of environmental conditions triggering or hinder-
ing fossilization (e.g., Arp et al. 2001). It is thus
Definition important to understand processes of fossilization
in order to identify the main factors controlling
Fossilization refers to the processes leading to the the formation of microfossils and better read the
preservation of traces of life in the geological paleontological record.
record. While metazoans and single-cell eukary- When a microorganism dies, organic mole-
otes leave undisputed traces in the geological cules and cellular structures are degraded very
record in the form of hard mineral parts, the rapidly. Enzymes released by dying cells and/or
fossilization of microorganisms such as Bacteria other cells catalyze this process (lysis). This
Fossilization, Process of 887
results in a rapid cycling of organic matter. Some disappearance, trichome disarticulation, varying
organic molecules might be selectively preserved terminal cell morphology, and rupture of sheath
because they are more resistant to this chemical (e.g., Toporski et al. 2002). If precipitation occurs
degradation such as polymers forming the cell in close connection with the microbial structures
wall of spores (e.g., sporopollenin) or molecules and if the minerals are small enough, very fine
transformed secondarily by chemical reactions cellular details can be preserved. For example, a
such as sulfurization of organic carbon (e.g., 40-nm-thick cell wall of Gram-negative bacteria
Lepot et al. 2009a; Damste et al. 1998) or forma- can be fossilized by calcium phosphates (e.g.,
tion of geopolymers (Vandenbroucke and Benzerara et al. 2004a) or by iron minerals (e.g.,
Largeau 2007). Precipitation of minerals on cells, Miot et al. 2009). Such a preservation of the
i.e., biomineralization, is another major process 40-nm-thick cell wall of Gram-negative bacteria
that allows fossilization of prokaryotes by has also been evidenced recently in phosphorite F
armoring cellular structures against the lytic action samples from Morocco dating from the Paleo-
of enzymes. Several mechanisms can favor min- cene (Cosmidis et al. 2013). Embryos of eukary-
eral precipitation within and/or at the surface of otes could be preserved as well in the laboratory
cells and usually involve specific metabolic activ- at different development stages by phosphatiza-
ities and environmental conditions. For example, tion, simulating what likely occurred during the
it has been proposed that anoxic conditions are fossilization of the famous ~630 Ma old
necessary for fossilization. The absence of O2 Doushantuo embryos (e.g., Yin et al. 2007).
might indeed limit aerobic respiration, hence deg- While some organic carbon might be degraded
radation or organic matter but is not a sufficient during these processes, it has been shown that
parameter (Allison 1988). In addition to this, most of it can be preserved within resulting min-
anoxic environments are usually rich in metals eralized microfossils (e.g., Benzerara
(e.g., Fe, Mn) and/or sulfides which can precipitate et al. 2004b). The diversity of organic moieties
on cells. that can be preserved in fossils still seems poorly
Experimental taphonomy refers to fossiliza- assessed. Biomarkers, i.e., highly resistant
tion experiments conducted in the laboratory or organic molecules comprising cell walls, are
in the field. Although these studies may not all obvious examples (e.g., Derenne et al. 2008).
reflect the full complexity of natural environmen- Some organic geochemistry studies have how-
tal conditions, they reveal biotic patterns and ever shown that additional molecules such as
fossilizable properties that have been overlooked proteins could be preserved more than a 100 mil-
so far. First, such studies which are still in their lion years (Riboulleau et al. 2002). Although
infancy have shown that fossilization of microor- viruses may have been major players in life evo-
ganisms can be achieved in few days or few lution (e.g., Forterre 2006), only little is known
weeks. For example, exposing bacterial cells to about their geological record. Interestingly, some
a solution rich in Ca, Fe, or Si can lead to their taphonomy experiments have been performed on
encrustation in few hours (Toporski et al. 2002; these “organisms.” Biomineralization experi-
Benzerara et al. 2004a; Ferris and Magalhaes ments on viruses show that dissolved iron can
2008; Miot et al. 2009). Moreover, these experi- penetrate virus capsids and bind to internal sites
ments help decipher what can be preserved in (Daughney et al. 2004). As a result, virus capsids
microfossils, regarding their chemistry as well can serve as nuclei for the growth of iron oxide
as their ultrastructure. Cyanobacteria have particles. The resulting morphology differs from
received particular attention. Intra- and extracel- abiotic iron oxides and organic molecules, which
lular silicification has been observed. It has been originally composed the capsids, can be effi-
shown that sheaths are preferentially preserved as ciently preserved by the iron minerals that
compared to walls and cytoplasm (e.g., Bartley armor them, increasing the possibility of accu-
1996). Some transformations could be observed rately identifying them (e.g., Kaiser and
including cell collapse, shrinkage and Guggenberger 2000).
888 Fossilization, Process of
After the precipitation of minerals on organic prebiotic chemistry experiments (e.g., Deamer
structures, further degradation of morphology et al. 2006), fluid inclusions, carbonaceous fila-
and organic molecules can take place. This mentous shapes resulting from migrating organic
stage is influenced by mechanical stresses, circu- matter (with carbon isotopic fractionation resem-
lation of fluids, and metamorphism, e.g., an bling life patterns) around minerals casts in
increase in temperature and pressure. Formation hydrothermal environments (Brasier 2005;
of carbonate nodules has been inferred as an Brasier et al. 2006), aggregates of silica spheres
efficient mechanism for fossil preservation (e.g., and rods in silica-rich waters of hydrothermal
Muller 1985). Observations of natural samples springs, migration of carbonaceous materials
have shown that microfossils can sometimes be along microfractures (VanZuilen et al. 2007),
preserved even after high-grade metamorphism within or around silica (e.g., Jones and Renaut
(e.g., Bernard et al. 2007), suggesting that T and 2007; Lepot et al. 2009b). Finally, mineralized
P might not be prominent factors in the disruption pseudo-fossils have been produced using a mix-
of microfossils. In contrast, mineral growth/ ture of barium carbonate and silica in laboratory
transformation has been shown to be highly dam- experiments (Garcia-Ruiz et al. 2003). The
aging (e.g., Oehler 1976). Metamorphic pro- resulting auto-assembling segmented filaments
cesses take place over much longer timescales, contain organic matter.
so it is more difficult to simulate them in the
laboratory. However, some recent studies pro-
vide an interesting way to address this issue, in Basic Methodology
particular, by noting that time and temperature
are inherently linked and that aging at low tem- Characterization of field samples using analytical
perature over long timescales can be simulated by tools at multiscale, including scanning electron
shorter aging at a higher temperature microscopy, transmission electron microscopy,
(Skrzypczak-Bonduelle et al. 2008). Raman spectromicroscopy, confocal laser
One key problem encountered in the study of scanning microscopy. Laboratory experiments
microfossils is that relatively complex morphol- simulating fossilization, diagenesis, and
ogies can be produced by purely abiotic processes metamorphism.
(e.g., Garcia-Ruiz et al. 2003) and can thus make
their identification difficult. Laboratory experi-
ments can be of value to identify possible specific Key Research Findings
features that might be used to discriminate abiotic
from biological objects. In addition to morphol- Fossilization is a very fast process. Fossilization
ogy, the composition of the organic matter pro- can preserve very fine details and chemical het-
duced by reactions such Fischer-Tropsch has erogeneities down to the nanometer-scale.
been carefully scrutinized and compared with
mature kerogens (e.g., McCollom and Seewald
2007). For example, a combination of infrared Applications
spectroscopy (providing an aliphaticity index of
the organic matter) and microscale measurements Search for ancient traces of life on Earth and
of the carbon isotopic compositions was used by elsewhere.
Sangely et al. (2007) to distinguish between biol-
ogy and Fischer-Tropsch-type reactions as
genetic processes for the bitumen found in the Future Directions
Cretaceous uranium deposits of Athabasca.
Known abiotic products that can mimic life mor- Several directions remain widely open: first, an
phologies or chemistries include vesicles made in improved characterization of microfossils in
the laboratory from meteoritic kerogen or in other ancient rocks which will allow in particular
Fossilization, Process of 889
discriminating between genuine microfossils and bacteria Ramlibacter tatahouinensis. Geomicrobiol

microfossil-like abiotic features. Most of the J 21:341–349
Benzerara K, Yoon TH, Tyliszczak T, Constantz B,
oldest traces of life are indeed still debated. The Spormann AM, Brown GE Jr (2004b) Scanning trans-
advance of cutting edge technologies, such as mission x-ray microscopy study of microbial calcifi-
Raman spectromicroscopy (Bernard et al. 2008; cation. Geobiology 2:249–259
Schopf and Kudryavtsev 2009), transmission Bernard S, Benzerara K, Beyssac O, Menguy N, Guyot F,
Brown GE Jr, Goffé B (2007) Exceptional preserva-
electron microscopy (e.g., Lepot et al. 2008), tion of fossil plant spores in high-pressure metamor-
focused ion beam milling (Wirth 2009), phic rocks. Earth Planet Sci Lett 262:257–272
synchrotron-based X-ray microscopy (e.g., Bernard S, Beyssac O, Benzerara K (2008) Raman map-
Lepot et al. 2009a; Obst et al. 2009), and ping using advanced line-scanning systems: geological
applications. Appl Spectrosc 62:1180–1188
NanoSIMS (e.g., Oehler et al. 2009), provides Brasier MD (2005) Critical testing of earth’s oldest puta-
analyses of mineral and organic matter down to tive fossil assemblage from the similar to 3.5 Ga Apex F
the nanoscale; this should help in achieving that chert, Chinaman creek, western Australia. Precam-
goal. Second, experimental taphonomy can be brian Res 140:55–102
Brasier M, McLoughlin N, Green O, Wacay D (2006)
developed further, inspired by studies of meta- A fresh look at the fossil evidence for early Archaean
zoans (e.g., Briggs and Kear 1993). A wide diver- cellular life. Phil Trans R Soc B 361:887–902
sity of biomineralizing microbial systems is now Briggs DEG, Kear AJ (1993) Fossilization of soft-tissue in
available and can provide pre-fossils. Their the laboratory. Science 259:1439–1442
Cosmidis J, Benzerara K, Gheerbrant E, Estève I,
extensive characterization down to the nanoscale Bouya B, Amaghzaz M (2013) Nanometer-scale char-
will offer new insight into what can be potentially acterization of exceptionally preserved bacterial fos-
preserved in the geological record. In addition, a sils in Paleocene phosphorites from Ouled Abdoun
significant improvement in the design of proto- (Morocco). Geobiology 11:139–153
Damste JSS, Kok MD, Koster J, Schouten S (1998)
cols to simulate diagenesis (in particular, the use Sulfurized carbohydrates: an important sedimentary
of fluids with an appropriate chemical composi- sink for organic carbon? Earth Planet Sci Lett
tion), aging, and metamorphism would comple- 164:7–13
ment this approach. Daughney CJ, Chatellier X, Chan A, Kenward P, Fortin D,
Suttle CA, Fowle DA (2004) Adsorption and precipi-
tation of iron from seawater on a marine bacteriophage
(PWH3A-P1). Mar Chem 91:101–115
See Also Deamer D, Singaram S, Rajamani S, Kompanichenko V,
Guggenheim S (2006) Self-assembly processes in the
prebiotic environment. Phil Trans R Soc
▶ Biomarkers, Morphological B 361:1809–1818
▶ Biomineralization Derenne S, Robert F, Skrzypczak-Bonduelle A,
▶ Bioprecipitation Gourier D, Binet L, Rouzaud JN (2008) Molecular
▶ Microfossils evidence for life in the 3.5 billion year old
Warrawoona chert. Earth Planet Sci Lett 272:476–480
Ferris FG, Magalhaes E (2008) Interfacial energetics of
bacterial silicification. Geomicrobiol J 25:333–337
References and Further Reading Forterre P (2006) The origin of viruses and their possible
roles in major evolutionary transitions. Virus Res
Allison PA (1988) The role of anoxia in the decay and 117:5–16
mineralization of proteinaceous macro-fossils. Paleo- Garcia-Ruiz JM, Hyde ST, Carnerup AM, Christy AG,
biology 14:139–154 Van Kranendonk MJ, Welham NJ (2003) Self-
Arp G, Reimer A, Reitner J (2001) Photosynthesis- assembled silica-carbonate structures and detection
induced biofilm calcification and calcium concentra- of ancient microfossils. Science 302:1194–1197
tions in phanerozoic oceans. Science 292:1701–1704 Grotzinger JP, Rothman DH (1996) An abiotic
Bartley JK (1996) Actualistic taphonomy of model for stromatolite morphogenesis. Nature
Cyanobacteria: implications for the Precambrian fossil 383:423–425
record. Palaios 11:571–586 Jones B, Renaut RW (2007) Microstructural changes
Benzerara K, Menguy N, Guyot F, De Luca G, Heulin T, accompanying the opal-A to opal-CT transition: new
Audrain C (2004a) Experimental colonization and evidence from the siliceous sinters of Geysir,
weathering of orthopyroxenes by the pleomorphic Haukadalur, Iceland. Sedimentology 54:921–948
890 Fossilized Microbial Mats
Kaiser K, Guggenberger G (2000) The role of DOM Skrzypczak-Bonduelle A, Binet L, Delpoux O, Vezin H,
sorption to mineral surfaces in the preservation of Derenne S, Robert F, Gourier D (2008) EPR of radicals
organic matter in soils. Org Geochem 31:711–725 in primitive organic matter: a tool for the search of
Lepot K, Benzerara K, Brown GE Jr, Philippot P (2008) biosignatures of the most ancient traces of life. Appl
Microbially influenced formation of 2, 724-million Magn Reson 33:371–397
year-old stromatolites. Nat Geosci 1:118–121 Toporski JKW, Steele A, Westall F, Thomas-Keprta KL,
Lepot K, Benzerara K, Brown GE Jr, Philippot P (2009a) McKay DS (2002) The simulated silicification of
Organic matter heterogeneity in 2.72 Ga stromatolites: bacteria – new clues to the modes and timing of bac-
alteration versus preservation by sulphur incorpora- terial preservation and implications for the search for
tion. Geochim Cosmochim Acta 73:6579–6599 extraterrestrial microfossils. Astrobiology 2:1–26
Lepot K, Philippot P, Benzerara K, Wang GY (2009b) van Zuilen M, Chaussidon M, Rollion-Bard C, Marty
Garnet-filled trails associated with carbonaceous mat- B (2007) Carbonaceous cherts of the Barberton Green-
ter mimicking microbial filaments in Archaean basalt. stone belt, South Africa: isotopic, chemical and struc-
Geobiology 7:1–10 tural characteristics of individual microstructures.
McCollom TM, Seewald JS (2007) Abiotic synthesis of Geochim Cosmochim Acta 71:655–669
organic compounds in deep-sea hydrothermal environ- Vandenbroucke M, Largeau C (2007) Kerogen origin,
ments. Chem Rev 107:382–401 evolution and structure. Org Geochem 38:719–833
Miot J, Benzerara K, Morin G, Kappler A, Bernard S, Wirth R (2009) Focused Ion Beam (FIB) combined with
Obst M, Férard C, Skouri-Panet F, Guigner JM, SEM and TEM: advanced analytical tools for studies
Posth N, Galvez M, Brown GE Jr, Guyot F (2009) of chemical composition, microstructure and crystal
Iron biomineralization by neutrophilic iron- structure in geomaterials on a nanometre scale. Chem
oxidizing bacteria. Geochim Cosmochim Acta Geol 261:217–229
73:696–711 Yin LM, Zhu MY, Knoll AH, Yuan XL, Zhang JM, Hu
Muller KJ (1985) Exceptional preservation in calcareous J (2007) Doushantuo embryos preserved inside dia-
nodules. Philos Trans R Soc Lond B Biol Sci pause egg cysts. Nature 446:661–663
311:67–73
Obst M, Wang J, Hitchcock AP (2009) Soft x-ray spectro-
tomography study of cyanobacterial biomineral nucle-
ation. Geobiology 7:577–591
Oehler JH (1976) Experimental studies in Precambrian Fossilized Microbial Mats
paleontology – structural and chemical changes in
blue-green-algae during simulated fossilization in syn-
▶ Stromatolites
thetic chert. Geol Soc Am Bull 87:117–129
Oehler DZ, Robert F, Walter MR, Sugitani K, Allwood A,
Meibom A, Mostefaoui S, Selo M, Thomen A, Gibson
EK (2009) NanoSIMS: insights to biogenicity and
syngeneity of Archaean carbonaceous structures. Pre-
cambrian Res 173:70–78
Foton Capsule, Spacecraft
Raff EC, Villinski JT, Turner FR, Donoghue PCJ, Raff RA
(2006) Experimental taphonomy shows the feasibility René Demets
of fossil embryos. Proc Natl Acad Sci U S A ESTEC (HSF-USL), Noordwijk,
103:5846–5851
The Netherlands
Riboulleau A, Mongenot T, Baudin F, Derenne S, Largeau
C (2002) Factors controlling the survival of proteina-
ceous material in late Tithonian kerogens (Kashpir oil
shales, Russia). Org Geochem 33:1127–1130 Definition
Riding R (2002) Structure and composition of organic
reefs and carbonate mud mounds: concepts and cate-
gories. Earth Sci Rev 58:163–231 Foton is an unmanned recoverable spacecraft,
Sangely L, Chaussidon M, Michels R, Brouand M, designed and manufactured by TsSKB-Progress
Cuney M, Huault V, Landais P (2007) Micrometer in Samara (Russia). Fifteen Fotons were launched
scale carbon isotopic study of bitumen associated
between 1985 and 2007, initially from Plesetsk,
with Athabasca uranium deposits: constraints on the
genetic relationship with petroleum source-rocks and later from Baikonur. The launch vehicle is a
the abiogenic origin hypothesis. Earth Planet Sci Lett Soyuz-U. The reentry capsule is spherically
258:378–396 shaped with a diameter of 2.3 m. Touchdown in
Schopf JW, Kudryavtsev AB (2009) Confocal laser scan-
the border zone of Russia and Kazakhstan. Pay-
ning microscopy and Raman imagery of ancient
microscopic fossils. Precambrian Res 173:39–49 load consisting of scientific and technological
Fractionation, Mass Independent and Dependent 891
experiments with often significant contributions chemical or physical properties. The chemical
from Western Europe. Power provided by batte- or isotopic composition of each phase may reflect
ries, flight duration therefore limited to an enrichment or depletion in one element or
10–16 days. Orbital parameters: inclination 63 , isotope with respect to the others, which can
period 90 min, orbital shape: near-circular, cruis- yield information about the mechanism of forma-
ing altitude around 300 km. Foton has frequently tion. Fractionation can be caused by differences
been used for space exposure studies in astrobi- in mass and binding energy and by biochemical
ology and astrochemistry (▶ Biopan) as well as reactions, which alter the ratio of one element or
reentry experiments (▶ Stone). Spacecrafts Mod- isotope to another. For example, fractionation of
ified Foton have been launched since the failure stable carbon isotopes can provide information
of Foton M1 in 2002 at launch. Since Foton M4, on the biotic or abiotic origin of some organic
which was successfully launched and retrieved in compounds, as photosynthetic reactions process F
2014, the spacecraft is equipped with solar panels the lighter isotope, 12C, more rapidly than 13C,
replacing former batteries. resulting in enrichment in 12C of its products.
See Also
See Also
▶ Biopan
▶ Stone ▶ Biomarker, Isotopic
▶ Deuterium/Hydrogen Ratio
▶ Differentiation, Planetary
▶ Distillation, Rayleigh
Fractional Abundances ▶ Fractionation, Mass Independent and
Dependent
▶ Molecular Abundances
▶ Isotope
▶ Isotopic Exchange Reaction
Fractionation ▶ Isotopic Fractionation (Planetary Process)
▶ Isotopic Ratio
Jennifer C. Stern ▶ Nitrogen Isotopes
Planetary Environments Laboratory, NASA
Goddard Space Flight Center, Greenbelt, MD,
USA
Fractionation, Mass Independent

Synonyms and Dependent
Differentiation; Disproportionation; Partitioning; Francis Albarède

Separation Ecole Normale Supérieure de Lyon, Lyon,
France
Definition
Keywords
▶ Fractionation is the partitioning of a chemical
species into two or more phases based on its Isotope; Mass fractionation
892 Fractionation, Mass Independent and Dependent
Definition 1. Pre-2.3 Ga sulfur also shows MIF of d33S with

respect to d34S.
Isotope fractionation is referred to as mass depen- Causes for solar system MIF are not agreed
dent when observed isotopic abundances deviate upon, at least not for all the elements. Self-
smoothly and monotonically with the masses of shielding calls for isotope-dependent shifts of
the isotopes from those in the reference material. absorption wavelengths in the UV range: dissoci-
Both kinetic and equilibrium processes are ation in the gas phase is proportional to the abun-
known to account for this most common form of dance of a particular isotope and not to its mass.
isotope fractionation. Alternate patterns are Self-shielding upon dissociation of a CO-rich gas
referred to as mass independent and are known is a popular interpretation for oxygen in the solar
for O and S in some specific environments and nebula, but critics invoke fast re-equilibration
are often related to photochemistry in the solar between the reaction products. Today, it is widely
nebula and in planetary atmospheres. believed that non-mass dependent fractionation
between the Sun and terrestrial planets is not
inherited from the pre-Solar constituents of the
Overview nebula. Symmetry is a critical property that defines
reaction paths: the two ozone isotopologues
Isotope fractionation varies approximately with (molecules that differ only in their isotopic com-
the difference of the isotopic masses. Using position) 16O 18O17O and 16O 18O16O have differ-
▶ oxygen isotopes as an example, the mass ent symmetries, and their reaction pathways
difference 18O 16O is +2 and d18O will there- involve different numbers of quantum states,
fore be twice the d17O where, e:g., d18 O is which greatly affects the rate of the reactions in
h i which ozone is involved. Ozone present in the
18
O=16 O sample = 18 O=16 O std 1 Þ:
modern stratosphere blocks solar UV radiation.
In a d17O versus d18O diagram, all terrestrial Prior to the “▶ Great Oxygenation Event” at
samples plot on the same fractionation line with a 2.3 Ga, the abundance of O2 in the atmosphere
slope of 0.5. Different planets plot on parallel was low and the ozone layer absent: photolysis by
fractionation lines, except the Earth and the solar UV of the tropospheric SO2 released by
Moon, which plot on the same line, a strong argu- volcanic activity is thought to have been an impor-
ment in favor of a common origin for the two tant cause of the MIF recorded in Archean sulfates.
planetary objects (the Moon Giant Impact). Finally, the finite volume of heavy nuclei and spin-
Small variations of the slope reflect different con- orbit coupling may be additional minor causes of
ditions of fractionation, notably kinetic and distil- deviation of isotope fractionation patterns from the
lation effects. Solar oxygen measured in the solar purely mass-dependent trends.
wind implanted in lunar metallic particles or in the
particles collected by the Genesis spacecraft con-
tains a mass-independent 16O excess of 5 percent.
See Also
Mass-independent isotope fractionation (MIF)
▶ Isotopic Fractionation (Planetary Process)
in the Solar System is observed for some ele-
▶ Oxygen Isotopes
ments, and the most noticeable effects occur for
▶ Sulfur Isotopes
O and S. The d17O and d18O of calcium-
aluminum-rich refractory inclusions (▶ CAIs)
and ▶ chondrules in ▶ chondrites tend to plot References and Further Reading
on a line with a slope of 1.0, instead of the normal
slope of 0.5 expected from mass-dependent frac- Sharp Z (2007) Principles of stable isotope geochemistry.
tionation. Likewise, the oxygen of some trace Prentice Hall, Upper Saddle River
Thiemens MH (2006) History and applications of mass-
atmospheric compounds, notably ozone, nitrate, independent isotope effects. Annu Rev Earth Planet
and sulfate, also plot on a line with a slope of Sci 34:217–262
Fragmentation of Interstellar Clouds 893
A starting point for many theories is the

Fragmentation of Interstellar Clouds observed fact that the parent cloud has far too
much mass, and is far too cold, to be supported by
Steven W. Stahler internal pressure. In the absence of other forces,
Department of Astronomy, University of the object must collapse on itself. The traditional
California, Berkeley, CA, USA view of fragmentation is that this collapse gener-
ates substructure. As the cloud contracts, it breaks
apart, producing several daughter clouds. Each
Keywords daughter, in turn, collapses and breaks up. After a
number of generations, fragments with stellar-
Star formation type mass are produced – the observed dense
cores. F
Within the last few decades, many theorists
Definition have performed numerical simulations of col-
lapsing clouds. The fragmentation hierarchy is
The dense cores that form stars through gravita- not seen. However, if the parent cloud is also
tional collapse are embedded in much larger and turbulent, it may directly create substructures
more rarefied expanses of gas. How the parent resembling dense cores. Intriguingly, the simu-
▶ molecular cloud produces its substructure of lated objects even exhibit the range of three-
dense cores is the problem of fragmentation. dimensional shapes of real cores.
The traditional view is that the parent cloud Unfortunately, there is little evidence that the
breaks apart as it collapses in on itself. In numer- larger bodies are undergoing collapse. Observa-
ical simulations, objects resembling dense cores tions indicate that the typical age of dark clouds
are created in turbulent, collapsing clouds. How- and clumps is 10 million years, an order of mag-
ever, there is little evidence that large clouds are nitude longer than the time for them to collapse. It
indeed collapsing. If they are not, but are at least thus appears that these entities are indeed
temporarily stable, then dense cores must be pro- supported, probably by their internal magnetic
duced in another fashion, perhaps by the slow fields and turbulent motion.
accretion of background gas. If the parent bodies are not collapsing, then
dense cores must arise in another way. Rather
than breaking off from the parent (the top-down
Overview view), they may accrete gas from their surround-
ings (the bottom-up view). The mass of the
The interstellar clouds that contain young stars ▶ dense core increases until the object becomes
are relatively small structures embedded within unstable and undergoes collapse. This picture is
more diffuse background gas. These dense cores in better accord with the observation that dense
have sizes of about 0.1 pc and masses comparable cores themselves appear to be in force balance, at
to that of the Sun. The diffuse parent bodies, least before they form stars.
known as dark clouds or clumps, have sizes larger
by two orders of magnitude and masses of at least
1,000 times solar. See Also
It is the gravitational collapse of dense
cores that creates the new stars we observe in ▶ Dense Core
them. But how are dense cores themselves cre- ▶ Interstellar Medium
ated? The general, and as yet unsolved, issue of ▶ Molecular Cloud
how these substructures arise within relatively ▶ Star Formation, Observations
large molecular clouds is the problem of ▶ Star Formation, Theory
fragmentation. ▶ Stellar Cluster
894 Free Amino Acid

Free Energy
Hoyle F (1953) On the fragmentation of gas clouds into
galaxies and stars. Astrophys J 118:513
Jacques Reisse
Offner S, Klein RL, McKee CF (2008) Driven and
decaying turbulence simulations of low-mass star for- Université Libre de Bruxelles, Brussels, Belgium
mation: from clumps to cores to protostars. Astrophys
J 686:1174
Stahler SW, Palla F (1994) Chapter 12: Multiple star
formation. In: The formation of stars. Wiley,
Definition
Weinheim
Free energy refers to the amount of energy in a
system that can be converted to work when the
reaction takes place at constant pressure. This
amount of energy is necessarily less than the
Free Amino Acid available enthalpy. For chemical reactions taking
place at constant pressure (which is frequently the
Henderson James (Jim) Cleaves II case), free energy refers to the Gibbs free energy.
Earth–Life Science Institute (ELSI), Tokyo The thermodynamic definition of the Gibbs
Institute of Technology, Meguro–ku, Tokyo, energy (G) leads to the familiar relation G = H
Japan TS, where H stands for enthalpy, T for temper-
Institute for Advanced Study, Princeton, NJ, ature, and S for entropy.
USA
Washington, DC, USA See Also
of Technology, Atlanta, GA, USA ▶ Bioenergetics
Definition
Free Radical
In chemistry, a free amino acid is an amino acid
which is not covalently bound in a peptide or in ▶ Radical
another sort of linkage, for example, in a
melanoidin polymer, an ▶ HCN polymer, or che-
lated to an inorganic ion.
Free Water
▶ Water Activity
See Also
▶ Amino Acid
▶ Amino Acid Precursors
▶ HCN Polymer Freefall
▶ Oligopeptide
▶ Protein ▶ Microgravity
Frost Line 895
Free-Fall Time FRET
Steven W. Stahler Henderson James (Jim) Cleaves II

Department of Astronomy, University of Earth–Life Science Institute (ELSI), Tokyo
California, Berkeley, CA, USA Institute of Technology, Meguro–ku, Tokyo, Japan
Definition Washington, DC, USA
The time required for an astronomical object to of Technology, Atlanta, GA, USA
collapse under the influence of self-gravity is F
called the free-fall time. A hypothetical gas
Synonyms
sphere of uniform density and zero temperature
collapses to a point at its center in a finite time.
Fluorescence resonance energy transfer; Förster
This time is inversely proportional to the square
resonance energy transfer
root of the initial density. The free-fall time of
real objects can still be approximated from the
idealized result. For stars, this time is roughly 1 h. Definition
Such objects are normally supported by internal
pressure and do not collapse. In this case, the FRET is a technique used to measure the proxim-
free-fall time is approximately equal to the ity of chemical groups in macromolecules or
sound-crossing time and to the period of global between host and guest molecules. It involves the
oscillations. use of a donor (in an excited electronic state) and
acceptor ▶ chromophore. When they are in close
proximity (usually <10 nm), the donor may trans-
fer its energy to the acceptor through nonradiative
See Also
dipole-dipole coupling. This is known as “Förster
resonance energy transfer.” When both chromo-
▶ Gravitational Collapse, Stellar
phores are fluorescent, the term “▶ fluorescence
▶ Fragmentation of Interstellar Clouds
resonance energy transfer” is used instead,
▶ Protostars
although the energy is not actually transferred by
fluorescence but by nonradiative transfer.
See Also
Free-Free Emission
▶ Chromophore
▶ Bremsstrahlung Radiation
▶ Fluorescence
French Space Agency Frost Line
▶ CNES ▶ Snow Line

896 Fullerane
Definition
Fullerane
A fullerene is a carbon molecule arranged in the
William M. Irvine form of a hollow sphere. Any graphene sheet can
University of Massachusetts, Amherst, MA, USA be closed into a fullerene cage provided that 12 pen-
tagons are inserted into the sheet of condensed
hexagonal rings. Only fullerenes having the 12 pen-
tagonal sites fully annealed by hexagonal rings are
Synonyms
stable and have been isolated in macroscopic quan-
tity. This is an important rule regulating the fuller-
Hydrogenated fullerene
ene stability. The fullerenes isolated in
macroscopic quantity are those following strictly
the isolated pentagons rule: C60, C70, C76, C84, C90,
Definition C94. . .. Each fullerene contains 2(10+z) carbon
atoms corresponding to 12 pentagonal sites and z
Fulleranes are hydrogenated derivatives of fuller- hexagons. This building principle is a consequence
enes. It has been speculated that they may be of Euler’s theorem (e.g., Kirk 2007). Fullerenes
present in the interstellar medium and in the can have elements (metals, noble gases) trapped
envelopes of certain types of stars. inside the cages, and these molecules are known as
endohedral fullerenes (Kroto et al. 1991).
See Also History
▶ Fullerene The structure of the most common fullerene, C60,

was first hypothesized in 1970 by Eiji Osawa in
Japan and was the object of topological and the-
oretical calculations by the Russians
I. V. Stankevich, D. A. Bochvar, and
Fullerene
E. G. Gal’pern in 1973. Only in 1984–1985,
using mass spectrometry, was it possible to
Franco Cataldo1,2 and Susana Iglesias-Groth3
1 observe a series of carbon clusters that were rec-
Istituto Nazionale di Astrofisica – Osservatorio
ognized as fullerenes. H. Kroto, R. Curl, and
Astrofisico di Catania, Catania, Italy
2 R. Smalley were awarded the 1996 Nobel Prize
Actinium Chemical Research, Rome, Italy
3 in Chemistry for their role in the discovery of this
Instituto de Astrofisica de Canarias, La Laguna,
new class of molecules. The production of fuller-
Tenerife, Spain
enes in microscopic quantities at first by
W. Kraetschmer and D. Huffmann in 1990 cre-
ated a major breakthrough in fullerene science.
Keywords
The fullerenes are named after Buckminster
Fuller, an architect who designed polyhedral
Carbon molecules; Hydrogenation; Spectros-
domes based on hexagonal and pentagonal faces.
copy; Interstellar molecules
Overview
Synonyms
Fullerene C60 is the most studied fullerene. It is
Buckminsterfullerene; Buckyball; Footballene composed of 60 sp2-hybridized carbon atoms
Fullerene 897
arranged in a truncated icosahedron geometry found. The main sources of carbon dust and mol-
resembling a soccer ball. The molecule contains ecules in the interstellar medium are the late-type
30 weakly conjugated double bonds located carbon-rich stars (Ehrenfreund and Charnley
between the hexagons. The diameter of C60 is 2000; Millar 2004), but there is a class of stars,
700 pm (picometers). All carbon atoms in C60 which lies in the transition between the asymp-
are equivalent, giving a sharp single signal in totic giant branch (AGB) and the ▶ planetary
the 13C-NMR spectrum located at a chemical nebula stage, that was thought as a very promis-
shift of 143.2 ppm (parts per million by fre- ing source of fullerenes. The stars in this class are
quency). The mean C-C distance is 141 pm, rather rare, are helium rich, and are extremely
which is almost same as in graphite (Kroto depleted in the hydrogen content of their gaseous
et al. 1991). shells, so that as the carbon vapor is ejected from
the star, it cools and forms an ideal environment F
for fullerene formation (Kroto 2006). In fact, the
Basic Methodology presence of hydrogen is known to have negative
effects on the formation of the fullerene cage and
Fullerenes are produced by quenching carbon to favor the production of ▶ polycyclic aromatic
vapor in a helium atmosphere. Carbon vapor hydrocarbons (PAHs) and other products (Goeres
can be generated by resistive heating of graphite and Sedlmayr 1992, 1993). The prototype of such
or in a 40–60-ampere carbon arc. The best yields stars where fullerenes may be present is
of fullerenes are obtained at a helium gas pressure R Coronae Borealis and the corresponding class
between 100 and 200 mbar. Under these condi- of RCrB stars (Unsold and Baschek 2002; Kaler
tions, a carbon soot is collected which, when 2006). Indeed, C60 fullerene has been found in
extracted with benzene or toluene, releases RCrB stars, but its detection was not easy and its
5–10 % of its weight under the form of extract- abundance low or negligible, in contrast with the
able colored matter consisting of a mixture of C60 expectations (Garcia-Hernandez et al. 2011).
and C70 fullerenes. C60 is by far more abundant On July 2010 the fullerenes C60 and C70 were
than C70 in the mixture. The two fullerenes can be found for the first time around a young ▶ plane-
separated by common chromatographic proce- tary nebula of our Galaxy known as Tc1 (Cami
dures on an alumina column using hexane/tolu- et al. 2010). The discovery was made possible
ene as eluents. Small amounts of higher through Spitzer infrared space telescope mea-
fullerenes (C76, C84, C90, C94) can be further surements. The detection of fullerenes in that
separated by high-performance liquid chroma- astrophysical object came as a surprise, since it
tography on reversed phase using acetonitrile/ is known that fullerenes are formed only in envi-
toluene as mobile phase. In the extracted soot, ronments completely free from hydrogen. The
fullerenes > C100 remain, which can be further fullerenes were detected in the inner core region
extracted with 1,2,4-trichlorobenzene (Taylor of Tc1, which turned out to be carbon rich, hydro-
1999). Fullerenes can be produced in relatively gen poor, and dusty (Cami et al. 2010). Further-
small yield also under controlled combustion more, from the spectral signature, it was evident
conditions in sooting flames and extracted with that C60 and C70 were embedded into the carbon
solvents from the recovered soot. This method dust and not in the vapor phase (Cami et al. 2010).
has achieved industrial applications (Murayama Since the discovery of Cami et al., especially
et al. 2004). fullerene C60 and sometimes also C70 have been
found in many other astrophysical objects, for
example, in planetary nebulae belonging to the
Key Research Findings Magellanic Clouds (Garcia-Hernandez
et al. 2010), in protoplanetary nebulae (Zhang
Fullerenes were predicted to be present in space and Kwok 2011), and subsequently in a number
(Hare and Kroto 1992) and have indeed been of planetary nebulae in our Galaxy and in the
898 Fullerene
Large and Small Magellanic Clouds (Garcı́a- medium and their theoretical spectral properties
Hernández et al. 2012). Fullerene C60 has been can be found in the recent works of Iglesias-Groth
also detected in the interstellar medium and more (2004, 2005, 2006). A connection between ful-
precisely in the reflection nebulae NGC 7023 and lerenes and the strongest ▶ diffuse interstellar
NGC 2023 (Sellgren et al. 2009, 2010). band (DIB) known in the optical has also been
In the interstellar medium, C60 fullerene proposed (Iglesias-Groth 2007, 2008).
should be present as a neutral molecule or may The reference infrared spectra of fullerenes
undergo ionization to C60+. This cation may be and fulleranes (the hydrogenated fullerenes) are
responsible for some spectral features in the dif- now available in a wide range of temperatures
fuse interstellar bands (DIBs), absorption bands useful for searching and recognizing such mole-
detected in the spectra of stars in our Galaxy, and cules (Iglesias-Groth et al. 2011, 2012). Further-
beyond (Foing and Ehrenfreund 1994). Further more, the molar extinction coefficients and
claims for the detection of C60+ in the ISM seem integrated molar absorptivity of these molecules
to confirm once again that this molecule is ubiq- are now available for the estimation of the abun-
uitous in space (Berné et al. 2013). Fullerene dance of fullerenes and fulleranes (Iglesias-Groth
cations may undergo multiple addition of atomic et al. 2011, 2012).
hydrogen, forming hydrogenated derivatives An important property of molecules of
(Petrie et al. 1995). Neutral fullerenes also add astrochemical interest regards their stability to
atomic hydrogen easily at very low temperatures UV photons. A plethora of organic molecules
(Howard 1993), so that it is reasonable to think are today known in different space environments
that fulleranes, the hydrogenated fullerene deriv- (Ehrenfreund and Charnley 2000), some of them
atives, should be present in the interstellar known to be the precursors of life. Photochemical
medium (Petrie and Bohme 2000; Cataldo and processing may lead to the production in space of
Iglesias-Groth 2010). other interesting molecules from common pre-
Fulleranes can be produced in the laboratory cursors or may lock certain molecules into dust
under a variety of conditions (Cataldo and or dust-forming nanoparticles. In this context, the
Iglesias-Groth 2010). Deuterated fulleranes photophysical and photochemical properties of
have also been synthesized. Significant isotope fullerenes and their hydrogenated counterparts,
effects both in the photolysis and in the thermal the fulleranes, are of interest, both for the detec-
decomposition of fulleranes and perdeutero- tion of these species and to estimate their survival
fulleranes have been observed (Cataldo 2009a; and their fate in the harsh space environment. The
Cataldo and Iglesias-Groth 2010). An enrichment photophysical properties of fullerenes are well
in the deuterium content in fulleranes is hence known, the photochemistry and the radiation
expected in the space environment. Additionally, chemistry of these molecules having been
fulleranes release molecular hydrogen under cer- explored, revealing a very high stability of fuller-
tain circumstances and are thought to play a role enes to both high-energy photons and corpuscular
in molecular hydrogen formation in space, radiation (Cataldo et al. 2009b) (Fig. 1).
starting from atomic hydrogen (Cataldo and
Iglesias-Groth 2010).
Calculated spectra for hydrogenated fuller- Future Directions
enes have been published in comparison with
the ▶ unidentified infrared emission bands The search for fullerenes in various natural envi-
(Webster 1991; Cataldo and Iglesias-Groth ronments has been the object of a monograph
2010). The infrared spectrum of C60H36 has also (Rietmeijer 2006). Fullerenes have been found
been compared with the infrared features of astro- in the Cretaceous-Tertiary boundary layer, in
physical objects like the protoplanetary nebulae meteorites, and in organic deposits. The search
(Cataldo 2003). An inventory about fullerenes for fullerenes in space will be intensified
and hydrogenated derivatives in the interstellar (Sellgren et al. 2009; Iglesias-Groth 2007, 2008;
Fullerene 899
fullerene which is completely soluble in water

and which can be considered a source of organic
carbon for the synthesis of simple molecules pre-
cursors of biochemistry (Iglesias-Groth
et al. 2013).
See Also
▶ Diffuse Interstellar Bands

▶ Fullerane
▶ Planetary Nebula F

Fullerene, Fig. 1 Stick and ball model of C60 fullerene
Berné O, Mulas G, Joblin C (2013) Interstellar C60+.
Astron Astrophys 550:L4, 5 pp
Lambert et al. 2001; Clayton et al. 1995; Goeres Cami J, Bernard-Salas J, Peeters E, Malek SE
and Sedlmayr 1993). It will be necessary to (2010) Detection of C60 and C70 in a young planetary
nebula. Science 329:1180–1182
record the laboratory spectra in the infrared, the Cataldo F (2003) Fullerane, the hydrogenated C60 fuller-
ultraviolet, and the visible of fullerenes and ene: properties and astrochemical considerations.
fulleranes, especially at low temperatures, in Fuller Nanotub Car Nanostruct 11:295–316
order to have available reference spectra useful Cataldo F, Iglesias-Groth S (2010) Fulleranes: the hydro-
genated fullerenes. Springer, Berlin
for the research of this class of molecules in Cataldo F, Iglesias-Groth S, Manchado A (2009a) On the
space. Moreover, future research in the labora- action of UV photons on hydrogenated
tory will be directed toward higher fullerenes, fulleranes C60H36 and C60D36. Roy Astron Soc
which until now have been scarcely studied sim- 400:291–298
ply because they are not easily accessible. Cataldo F, Strazzulla A, Iglesias-Groth S (2009b) Stability
of C60 and C70 fullerenes toward corpuscular and
Fullerenes have been found mixed with PAHs
gamma radiation. Roy Astron Soc 394:615–623
in certain astrophysical environments, and it is Cataldo F, Garcı́a-Hernández DA, Manchado A (2013)
completely reasonable that both fullerenes and Sonochemical of fullerene C60/anthracene Diels-
PAHs share the same destiny when ejected into Alder mono and bis-adduct. Fuller Nanotub Car
Nanostruct 21 (in press)
the interstellar medium. One interesting aspect of
Clayton GC, Kelly DM, Lacy JH, Little-Marenin IR,
future research regards the reactivity of fullerenes Feldman PA, Bernath PF (1995) A mid-infrared search
with PAHs. We know already that fullerenes for C60 in R coronae borealis stars and IRC + 10216.
easily merge with linear PAHs to form Diels- Astronom J 109:2096
Ehrenfreund P, Charnley SB (2000) Organic molecules in
Alder adducts known as acenes (Cataldo
the interstellar medium, comets and meteorites: a voy-
et al. 2013). The spectra of adducts of fullerenes age from dark clouds to the early Earth. Annu Rev
and PAHs prepared in the lab could be used as Astronom Astrophys 38:427–483
reference when searching for evidence of the Foing BH, Ehrenfreund P (1994) Detection of two inter-
stellar absorption bands coincident with spectral fea-
reactivity of fullerenes with PAHs in space.
tures of C 60 +. Nature 369:296–298
Another intriguing aspect regards the fact that Garcı́a-Hernández DA, Manchado A, Garcı́a-Lario P,
fullerenes can be a source of carbon for prebiotic Stanghellini L, Villaver E, Shaw RA, Szczerba R,
synthesis on icy grains or under other conditions. Perea-Calderón JV (2010) Formation of fullerenes in
H-containing planetary nebulae. Astrophys J Lett 724:
Fullerene C60 is insoluble in water, but if
L39–L43
radiolyzed in a water medium or in water- Garcı́a-Hernández DA, Kameswara Rao N, Lambert DL
ammonia medium, it yields a polyhydroxylated (2011) Are C60 molecules detectable in circumstellar
900 Fulminic Acid
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Garcı́a-Hernández DA, Villaver E, Garcı́a-Lario P, perspectives. Kluwer, Dordrecht, pp 17–31
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Shaw RA, Cataldo F (2012) Infrared study of fullerene (2004) Fullerene production in tons and more: from
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events due to dust formation. Astronom Astrophys ion/molecule reactions of fullerenes: chemical deriva-
265:216–236 tization of fullerenes within dense interstellar clouds
Goeres A, Sedlmayr E (1993) Hydrogen-blocking in C60 and circumstellar shells. Astrophys J 540:869–885
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25:106–112 Rietmeijer FJH (2006) Natural fullerenes and related struc-
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fulleranes and galactic anomalous microwave emis- ton TM, Joblin C (2010) C60 in reflection nebulae.
sion. Astrophys J 632:L25–L28 Astrophys J Lett 722:L54–L57
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Iglesias-Groth S, Hafez Y, Angelini G, Cataldo F (2013) g ▶ HCNO Isomers
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doi:10.1007/s10967-013-2484-0
Kaler JB (2006) The Cambridge encyclopedia of stars.
Fumarole
Cambridge University Press, Cambridge, p 203
Kirk A (2007) Euler’s polyhedron formula. Plus Mag Jörn Helbert
Kroto HW (2006) Introduction: space-Pandora’s box. In: DLR, Institut f€ur Planetenforschung, Berlin,
Rietmeijer FJH (ed) Natural fullerenes and related
Germany
structures of elemental carbon. Springer, Dordrecht,
pp 1–5
Kroto HW, Allaf AW, Balm SP (1991) C60: Buckminster-
fullerene. Chem Rev 91:1213–1235 Definition
Lambert DL, Rao NK, Pandey G, Ivans II (2001) Infrared
space observatory spectra of R Coronae Borealis stars.
I. Emission features in the interval 3–25 microns. Fumaroles (Latin fumus, smoke) are vents from
Astrophys J 555:925–931 which volcanic gas escapes into the atmosphere.
Fungi 901
Fumaroles may occur along tiny cracks or long Definition

fissures, in chaotic clusters or fields, and on the
surfaces of lava flows and thick deposits of pyro- Fungi are nonphototrophic, heterotrophic eukary-
clastic (explosive ash) flows. They may persist otic microorganisms that contain rigid cell walls
for decades or centuries if they are above a per- and produce spores. Fungi form a tight phyloge-
sistent heat source or disappear within weeks to netic cluster.
months if they occur atop a fresh volcanic deposit
that quickly cools.
Fumaroles are common on ▶ Earth; for exam- Overview
ple, there are an estimated four thousand fuma-
roles within the boundaries of Yellowstone Defining fungi within the group of eukaryotes is a
National Park. The Mars Exploration Rover complex and difficult task and it is preferable to F
Spirit has identified possible fumarolic deposits construct a definition based on their common
in Gusev crater. properties rather than on the differences that sep-
arate them from the rest of the eukaryotic species.
Their peculiar method of reproduction is perhaps
one of the main axis on which it is possible to
See Also construct a phylogenetic classification, reflecting,
in some way, the mechanisms of evolution of
▶ Crater, Impact each of the groups of fungi represented on Earth.
▶ Cryovolcanism Many species behave according to Mayr’s
▶ Earth species definition, presenting a sexual life cycle
▶ Mars where two mycelia having sexually compatible
cells join to produce a panmictic population that
mates randomly after meiotic division generating
a fertile population, while others are parasexual
(i.e., involving nuclear fusion followed by grad-
ual de-diploidization).
Functional Inhibitor In a high percentage of species, this mecha-
nism implies an asexual reproduction system that
▶ Antibiotic gives rise to specialized cells known as conidio-
phores that produce dikaryotic cells by mitosis
(only nuclear division) which are able to generate
a new organism. When they are isolated in pure
culture, the asexual strain, known as
Fungi anamorphous (i.e., Aspergillus nidulans), is
obtained by the mating of two anamorphous,
Aldo González while the sexual strain is known as
Centro de Biologı́a Molecular, CBMSO Consejo teleomorphous (i.e., Emericella nidulans). One
Superior de Investigaciones Cientificas species has these two reproductive states
Universidad Autónoma de Madrid, Madrid, (asexual and sexual). These two ways of repre-
Spain sentation in the world of living beings make it
necessary to create two systems of classification,
one for each type of reproductive state, knowing
Keywords beforehand that the one for the sexual cycle is
close to a phylogenetic proposal and that the
Aspergillus; Carpophores; Filamentous fungi; other, based on the forms of conidia production,
Hyphae; Mushrooms; Penicillium; Yeasts is rather artificial.
902 Fungi
About 120,000 species of fungi have been and plants. (11) They have a cosmopolitan distri-
described to date, although the total estimated bution. (12) Degradation, they are important
number of species is around 1.5 million. The ver- degraders of plant eliminate macro-polymers (i.e.,
satility of their “soma,” also named mycelia, cellulose, lignin); of fossil combustibles; “seques-
which in the case of filamentous fungi are made ters” of heavy metals and radioactive elements at
up of a group of septate hyphae, allow them to be high concentrations.
present in all possible ecosystems on Earth. Where In general they are responsible for 75 % of the
they are a minority, it is because this environment turnover of carbon (▶ Carbon Cycle), contribut-
has, in fact, been little studied yet. Recent studies ing to the degradation of biomass and facilitating its
of metagenomics report their presence in ecosys- reincorporation into the vital cycle of terrestrial and
tems under environmental pressure or in the sea. In aquatic plants. At present, the objective is to pro-
recent studies of classification of all described mote a new taxonomy based directly on compari-
species, six kingdoms have been established. sons of selected DNA sequences that encode genes
Fungi have been included in the kingdom Mycota, with a conserved biological function, instead of or
in which seven monophyletic “true fungi” are in addition to phenotypic characteristics. These
included: Chytridiomycota, Zygomycota, gene sequences should allow the construction of
Endomycota (Yeasts), Ustomycota, Ascomycota, phylogenetic trees integrating microscopic, ultra-
Basidiomycota, and Deuteromycota, all grouped structural, and biochemical data leading to a fuller
as Eumycota. The organisms known as understanding of fungal taxonomy with monophy-
“pseudofungi” are phylogenetically close to Pro- letic criteria. Nevertheless, the most reliable current
tists and contain the Oomycota and criterion to define a given species continues to be
Hyphochytridio-Labrynthulomycota groups. the cross-linking of two sexually compatible
According to their special way of life, all fungi monokaryon mycelia to form dikaryotic mycelia
can be defined by the following characteristics: able, in turn, to produce carpophores.
(1) They are heterotrophs and their nutrition is by
absorption. (2) In the vegetative stage, they grow
on or inside the substrate as hyphae forming a See Also
mycelium showing internal protoplasmic stream-
ing. Reproductive stages with mobility may occur. ▶ Carbon Cycle, Biological
(3) Cell wall is generally formed by glucans and ▶ Eukarya
chitin and, in some cases, glucans and cellulose. ▶ Eukaryote
(4) Nuclear status, Eucariota, uni- or multinucleate, ▶ Heterotroph
the thallus may be homo- or heterokaryotic, ▶ Phylogeny
dikaryotic or diploid, the last usually of short dura- ▶ Yeast
tion with exceptions in some groups. (5) Their life
cycle is simple although in some cases complex.
(6) Propagules are typical little spores produced in
high amounts. (7) Sporocarps or carpophores can Barnett JA, Payne RW, Yarrow W (1983) Yeasts: charac-
be microscopic or macroscopic, with characteristic teristics and identification. Cambridge University
shapes. In some groups they are known as mush- Press, Cambridge
room and are limited to differentiation tissues. Boddy L, Coleman M (2010) From another kingdom – the
amazing world of fungi. Royal Botanical Garden,
(8) Habitat, ubiquitous in terrestrial, freshwater Edinburgh
habitats and in lower numbers in marine waters, Cavalier-Smith T (1998) A revised six-kingdom system of
although fungal species have been less studied in life. Biol Rev Camb Philos Soc 73:203–266
this environment. (9) In ecology, they play impor- Cavalier-Smith T (2001) What are fungi? In: McLaughlin
DJ, McLaughlin EG, Lemke PM (eds) The mycota VII
tant roles as saprotrophs, mutualistic symbionts, A: systematics and evolution. Springer, Berlin, pp 3–37
parasites, or hyperparasites. (10) In pathology, Crous PW, Samson RA, Gams W, Summerbell RC,
they are the cause of serious pathologies to animals Boekhout T, de Hoog GS, Stalpers JA (2004a) CBS
Furanose 903
centenary: 100 years of fungal biodiversity and ecol- Definition

ogy. Stud Mycol 50(1):1–298
Crous PW, Samson RA, Gams W, Summerbell RC,
Boekhout T, de Hoog GS, Stalpers JA (2004b) CBS The term furanose denotes a five-component
centenary: 100 years of fungal biodiversity and ecol- cyclic structure containing four carbon atoms
ogy. Stud Mycol 50(2):299–586 and one oxygen atom and is generally reserved
de Hoog GS (2005) Fungi of the Antarctic: evolution for the description of the molecular framework in
under extreme conditions. Stud Mycol 51:1–79
Ellis MB (1976) Dematiaceous hyphomycetes. CMI, Kew monosaccharides. The furanose ring can be a
Emmons CW, Chapman YHB, UTZ JP, Kwon-Chung KJ cyclic hemiacetal of an aldopentose (e.g., ribose
(1977) Medical mycology, 3rd edn. Lea & Febiger, in RNA and deoxyribose in DNA) or a cyclic
Philadelphia hemiketal of a ketohexose (e.g., fructose).
Kendrick B (ed) (1992) The fifth kingdom. Focus Publish-
ing, R. Pullins Co, Newburyport
Kinghorn JR, Turner G (1992) Applied molecular genetics F
of filamentous fungi. Blackie Academic & Profes- Overview
sional, Glasgow
Kirk PM, Cannon PF, Minter DW, Stalpers JA
(2008) Ainsworth and Bisby’s dictionary of the Furanose denotes a five-membered cyclic struc-
fungi, 10th edn. CABI, Wallingford ture, belonging to a class of compounds termed
Kreger-van Rij NJW (ed) (1984) The yeasts, a taxonomy carbohydrates, which are molecules composed of
study. Elsevier Science Pub. B.V., Amsterdam carbon, oxygen, and hydrogen with the empirical
Ryvarden L (1991) Genera of polypores. Nomenclature
and taxonomy. Synopsis fungorum 5 fungiflora. formula Cn(H2O)n. Furanoses are differentiated
Gronlands Grafiske A/S, Norway by various means, including the carbon frame-
Sutton BC (1980) The coelomycetes. Fungi imperfecti work from which hydroxyl groups are attached,
with pycnidia, acervuli and stromata. CMI, Kew the stereochemistry of the hydroxyl groups at
Tavares II (1985) Laboulbeniales (fungi, ascomycta).
Micologia/memoir, vol 9. J Cramer Publisher, each carbon (R or S), and the conformation of
Braunschweig the overall structure (envelope or twist). Fura-
Von Arx JA (1981) The genera of fungi sporulating in noses are either assigned a D- or
pure culture. J Cramer, Germany L-configuration depending on the ▶ chirality of
Webster J, Weber R (2007) Introduction to fungi, 3rd edn.
Cambridge University Press, Cambridge the stereocenter at the carbon farthest away from
Whittaker RH (1969) New concepts of kingdoms of the aldehyde.
organisms. Science 163:150–160 Furanose form can exist in an equilibrium
Zycha H, Siepmann R, Linneman G (1969) Mucorales. distribution between a cyclic hemiacetal or cyclic
Eine beschreibung aller gattungen und arten dieser
pilzgruppe. Verlag von J Cramer, M€ unchen hemiketal and an uncyclized free ▶ aldehyde or
ketone, respectively. Entropy effects favor the
intramolecular reaction of the aldehyde
(or ketone) and hydroxyl group leading to the
Furanose distribution of forms as: hemiacetal >99.99 %,
free aldehyde ~0.01 % and the hydrate in trace
Ramanarayanan Krishnamurthy, amounts (Drew et al. 1998). The two predomi-
Tammy Campbell and Eun-Kyong Kim nant hemiacetals formed are furanose and
Chemistry, The Scripps Research Institute, ▶ pyranose. In general, furanoses are kinetically
La Jolla, CA, USA preferred, but thermodynamically less stable than
pyranoses. The anomeric carbon produced during
cyclic hemiacetal formation is bonded to two
Keywords oxygens. One oxygen is located inside the ring
and the other as a hydroxyl substituent in either
Alpha- vs beta-configuration; Carbohydrates; endo (beta)- or exo (alpha)-position relative to
Monosaccharide; Five-membered ring; D- or the ring. These are diastereomers or anomers of
L-chirality; Anomeric carbon; Conformation; furanose and generally have altered ratios due to
Aldehyde; Alcohol epimerization in solution.
904 Fusion Crust
The variations between the aldehydes, pyra- References and Further Reading
noses, and furanoses are biologically important
since nature favors one structure over the other. Drew KN, Zajicek J, Bondo G, Bose B, Serianni AS
(1998) 13C-labeled aldopentoses: detection and quan-
In the case of aldopentoses, four monosaccha-
titation of cyclic and acyclic form by heteronuclear 1D
rides exist: arabinose, ▶ ribose, xylose, and and 2D NMR spectroscopy. Carbohydr Res
lyxose. The equilibrium formation of furanose 307:199–209
from each aldopentose varies in solution Eschenmoser A (1999) Chemical etiology of nucleic acid
structure. Science 284:2118–2124
according to their stereogenic centers. Important
Eschenmoser A, Dobler M (1992) Why pentose and not
keto-furanoses in biology include ribulose and hexose nucleic acids? Helv Chim Acta 75:218–259
xylulose. Ribulose is converted to ribose in the Pitsch S, Wendeborn S, Krishnamurthy R, Eschenmoser
pentose phosphate pathway which is then used by A (2003) Pentopyranosyl oligonucleotide systems. 9th
communication. Helv Chim Acta 86:4270–4363
cells in the synthesis of nucleoside triphosphates
Prieur B (2001) Étude de l’Activité Prébiotique
and ▶ nucleic acids. Potentielle de l’Acide Borique. CR Acad Sci Ser II
The molecular basis of why ▶ RNA is C Chem 4:667–670
founded on a furanose and not a pyranose struc- Ricardo A, Carrigan M, Olcott N, Benner S (2004) Borate
minerals stabilize ribose. Science 303:196
ture (Eschenmoser 1999) is a key question that
Sacerdote MG, Szostak JW (2005) Semipermeable lipid
has been addressed extensively in the field bilayers exhibit diastereoselectivity favoring ribose.
concerning the origins of life. One study revealed Proc Natl Acad Sci U S A 102:6004–6008
limitations of the conformational diversity of
canonical nucleic acids by systematically synthe-
sizing alternative nucleic acid structures. Mark-
edly lower level of helicity was exhibited by
Fusion Crust
pentopyranosyl nucleic acids in comparison to
Tilman Spohn
ribonucleic acid, lending support to the idea that
the pentofuranose structure is responsible for the
helical shape of the duplex in RNA and ▶ DNA
Germany
(Eschenmoser and Dobler 1992; Pitsch
et al. 2003). Another study explored the assem-
blage and accumulation of pentofuranoses through
Definition
a borate-mediated stabilization of cis-diols in fura-
nose form (Prieur 2001; Ricardo et al. 2004). Inter-
A fusion crust is a feature of the external appear-
estingly, among the four aldopentoses, ribose has
ance of ▶ meteorites. It describes a glassy coating
the highest percentage of furanose form (Drew
of the meteorite. The crust forms as a conse-
et al. 1998). Furthermore, model protocellular
quence of the frictional heating and melting of
membranes exhibit a diastereoselective advantage
the most external layer of the meteorite as it
for ribose in the uptake of carbohydrates from
passes through the atmosphere and descends to
solution (Sacerdote and Szostak 2005). Together,
the ▶ Earth’s surface. Most of the melt is lost due
these findings suggest a reason for why ribose
to ▶ ablation. Thus, the fusion crust is mostly
(in its furanosyl form) may have been selected as
about 1 mm thick.
the structural basis upon which modern biology is
based.
See Also
See Also
▶ Ablation
▶ Monosaccharide ▶ Earth
▶ Ribose ▶ Meteorites
G
Ga See Also
Nicholas Arndt ▶ Earth, Age of

ISTerre, Université Grenoble Alpes, France ▶ Geochronology
▶ Ma
Synonyms
Giga-annum; Gigayear; Gyr Gabbro
Daniele L. Pinti
Definition
Ga is a common scientific abbreviation for
gigayears, 109 years, derived from the Latin
giga-annum. Note that the Latin accusative,
annum, expresses an absolute age, while the Definition
English accusative, years, expresses a
period of time. It is widely used in fields such as Gabbro is a common mafic intrusive magmatic
astronomy and geology. For example, the rock, chemically equivalent to the ▶ basalt.
current best estimate for the age of the universe, A medium to coarse-grained, dark-colored rock,
from the Wilkinson Microwave Anisotropy gabbro is composed of Ca-plagioclase and
Probe (spacecraft) data on the cosmic microwave clinopyroxene, with or without ▶ olivine and
background radiation, is 13.7 Ga, and the age of orthopyroxene. Earth’s oceans are underlain by
the Earth, commonly taken as the time of the gabbro which comprises a 3–5 km-thick layer of
completion of accretion, is 4.56 Ga. In geology, the ▶ oceanic crust, produced by crystallization
the term is often used to signify time before the of basaltic magma erupting at the ▶ Mid-ocean
present; for example, the time of the Moon- Ridges. Gabbro, and its extrusive equivalent
forming impact is 4.51 Ga, and the age of the basalt, is common on the ▶ Moon, ▶ Mars,
oldest rocks at the Earth’s surface is about many large asteroids, and probably ▶ Mercury
4.03 Ga. and ▶ Venus.

DOI 10.1007/978-3-662-44185-5
906 Gaia Hypothesis
See Also composition suggesting that life on Earth func-

tions as a homeostatic feedback system. He called
▶ Basalt this self-regulating living system Gaia, after the
▶ Oceanic Crust Greek goddess. Since 1971, the distinguished
▶ Ophiolite biologist Lynn Margulis collaborated closely
▶ Mid-Ocean Ridges with Lovelock in developing the Gaia Hypothe-
sis, and scientific experiments have provided a
number of useful predictions to support the
hypothesis, although it has always had highly
scientifically respected detractors. After long
Gaia Hypothesis
debate and much criticism, a modified Gaia
Hypothesis has found acceptance in the field of
Ricardo Amils
ecology. Ecologists generally consider the bio-
sphere an ecosystem, and the Gaia Hypothesis is
Universidad Autónoma de Madrid, Madrid,
consistent with today’s vision of global ecology,
Spain
which takes into consideration the interactions
between biota, the oceans, the geosphere, and
the atmosphere.
Definition
According to the Gaia Hypothesis, the biosphere See Also

and the physical components of the Earth form
a complex interacting system that maintains ▶ Autopoiesis
the climatic and biogeochemical Earth conditions ▶ Homeostasis
in homeorhesis. It was originally proposed by
James Lovelock who called it the Earth References and Further Reading
feedback hypothesis, and it is frequently
described as a way of seeing the Earth as a single Lovelock JE (1965) A physical basis for life detection
organism. experiments. Nature 207(7):568–570. doi:10.1038/
207568a0
Lovelock JE (1972) Gaia as seen through the atmosphere.
Atmos Environ 6(8):579–580. doi:10.1016/0004-
6981(72)90076-5
Overview Lovelock JE (1995) The ages of Gaia: a biography of
our living earth. Norton, New York. ISBN
James Lovelock first formulated the Gaia 0-393-31239-9
Hypothesis in the 1960s, as a result of his work Lovelock JE (2000) Gaia: a new look at life on
earth. Oxford University Press, Oxford. ISBN
for NASA on developing methods of detecting 0-19-286218-9
life on Mars. At first, the theory was a way to Lovelock JE, Margulis L (1974) Atmospheric homeostasis
explain the stable concentrations of chemicals by and for the biosphere- the Gaia hypothesis. Tellus
such as oxygen and methane that persisted in 26(1):2–10
Lovelock JE (1990) Hands up for the Gaia hypothesis.
Earth’s atmosphere. Lovelock suggested that
Nature 344:100–102. doi:10.1038/344100a0
detecting such unstable combinations in other Margulis L (1999) Symbiotic planet: a new look at evolu-
planets’ atmospheres was a relatively reliable tion. Basic Book, Houston
and cheap way to detect life. Schwartzman D (2002) Life, temperature, and the earth:
the self-organizing biosphere. Columbia University
Lovelock tried to explain the existence of a
Press, New York. ISBN 0231102135
global control system over surface temperature, Volk T (2003) Gaia’s body: toward a physiology of earth.
the salinity of the oceans, and the atmospheric MIT Press, Cambridge. ISBN 0-262-72042-6
Gaia Mission 907
satellite to reach its operational environment on a

Gaia Mission Lissajous orbit at Sun-Earth ▶ Lagrangian point
L2, 1.5 million kilometers away from Earth. At
Alessandro Sozzetti the time of this writing, commissioning opera-
Istituto Nazionale di Astrofisica tions are ongoing. During its 5 years of opera-
(INAF) – Osservatorio Astrofisico di Torino, tional lifetime (with the possible extension of an
Pino Torinese, Italy extra year), Gaia will monitor all point sources in
the visual magnitude range V = 6–20 mag, a
huge database of 109 stars, a few million galax-
Keywords ies, half a million quasars, and a few hundred
thousand asteroids.
Astrometry; Exoplanet; Extrasolar planet; Plane- As for the observing strategy, Gaia’s mode of
tary systems operation has adopted the principles successfully
experimented with the Hipparcos mission (ESA G
1997). In particular, it will continuously scan the
Definition sky, implying that all detected objects,
irrespective of their magnitudes, are observed
The ▶ European Space Agency’s Gaia all-sky for the same amount of time during each
survey will monitor astrometrically, during its field-of-view crossing, with mission-end
5-year nominal mission lifetime, all point sources observing time mainly depending on ecliptic lat-
(▶ stars, ▶ asteroids, quasars, extragalactic itude (Lindegren 2010). In this way, it is antici-
▶ supernovae, etc.) in the visual ▶ magnitude pated that Gaia will determine the five basic
range V = 6–20 mag, a huge database astrometric parameters (two positional coordi-
encompassing 109 objects. It was launched suc- nates, two proper motion components, and the
cessfully in December 2013. Using the continu- parallax) for all objects, with end-of-mission
ous scanning principle first adopted for (sky-averaged) precision between 7 and 25 m as
▶ Hipparcos, Gaia will determine the five basic (microarcseconds) down to the Gaia magnitude
astrometric parameters (two components of posi- G = 15 mag and a few hundred mas at
tion, two of ▶ proper motion, and the ▶ parallax) G = 20 mag, depending on color. Red objects
for all objects, with end-of-mission precision are expected to have better astrometry, while
exceeding that of ▶ Hipparcos by 1–2 orders of that for extremely blue targets is estimated to
magnitude. Gaia astrometry, complemented by degrade by a factor of 2.
onboard spectrophotometry and (partial) radial- The main partners inside the Gaia project are
velocity information, will have the precision nec- (1) the European Space Agency (ESA), which has
essary to quantify the early formation and subse- the overall project responsibility for funding and
quent dynamical, chemical, and star formation procurement of the satellite, launch, and opera-
evolution of the Milky Way Galaxy. tions; of interest is the fact that in this case,
satellite procurement includes the payload and
its scientific instruments, unlike ESA’s other sci-
Overview ence missions for which scientific instruments are
usually principal investigator led and funded (or,
The Gaia mission is the new global, all-sky, at least, co-funded) by participating national
astrometric initiative of the European Space space agencies; (2) European Aeronautic
Agency (ESA). Its launch occurred perfectly on Defence and Space Company Astrium, which
schedule from the ESA’s Kourou site in the was selected in 2006 as the prime industrial con-
French Guiana on December 19, 2013. tractor for designing and building the satellite
A Soyuz-Fregat launcher took the Gaia module according to the scientific and technical require-
to a transfer orbit, which in 1 month allowed the ments formulated to fulfill the mission science
908 Gaia Mission
case as approved at time of selection (ESA 2000); A combination of an ambitious science case,
and (3) the Gaia Data Processing and Analysis wishing to address breakthrough problems in
Consortium (DPAC), charged with designing, Milky Way astronomy, and lessons learned from
implementing, and running a complete software the Hipparcos experience brought European
system for the scientific processing of the satellite astronomers to realize that Gaia astrometry needed
data, resulting in the edition of the “Gaia Cata- to be complemented by onboard spectrophotome-
logue” a few years after the end of the operational try and (only for objects brighter than G = 17)
(observation) phase. radial-velocity information. These data will have
DPAC was formed in 2006 in response to an the precision necessary to quantify the early for-
“Announcement of Opportunity” issued by ESA. mation and subsequent dynamical, chemical, and
The consortium lists nearly 400 individual mem- star formation evolution of our Galaxy. The broad
bers in more than 20 countries. Six data range of crucial issues in astrophysics that will be
processing centers participate in the activities of addressed by the wealth of the Gaia data is sum-
the consortium, which is organized in eight marized by, e.g., Perryman et al. (2001). One of the
“Coordination Units,” each responsible for the relevant areas on which the Gaia observations will
development of one part of the software (e.g., have great impact is the astrophysics of planetary
core astrometric processing, photometry). The systems (e.g., Casertano et al. 2008), in particular
ninth coordination unit (CU9), the last DPAC when seen as a complement to other techniques for
coordination unit, will be in charge of the design exoplanet detection and characterization (e.g.,
and implementation of the Gaia archive. CU9 Sozzetti 2010).
was created separately from the rest of DPAC
through a specific Announcement of Opportunity
by ESA. Most of the financial support is provided Basic Methodology
by ESA and by the various national space agen-
cies through a legally binding long-term funding The problem of the correct determination of the
agreement, a real first for ESA-run missions. astrometric orbits of planetary systems using
There will be no proprietary periods for the Gaia data (highly nonlinear orbital fitting proce-
scientific exploitation of the data. The final Gaia dures, with a large number of model parameters)
Catalogue will be produced and immediately will present many difficulties. For example, it
delivered to the astronomical community world- will be necessary to assess the relative robustness
wide as soon as ESA and DPAC will agree on the and reliability of different procedures for orbital
processed data having reached the targeted fits, together with a detailed understanding of the
(science) quality. This catalogue is expected to statistical properties of the uncertainties associ-
be ready 3 years after the end of operations. ated with the model parameters. For multiple
Finally, an intermediate data release scenario systems, a trade-off will have to be found
has been established, with a preliminary schedule between accuracy in the determination of the
of four successive intermediate releases (the first mutual inclination angles between pairs of plan-
occurring approximately 2 years after launch) of etary orbits, single-measurement precision, and
increasingly more complete astrometric, photo- redundancy in the number of observations with
metric, and spectroscopic data. Photometric sci- respect to the number of estimated model param-
ence alerts will instead be produced and released eters. It will constitute a challenge to correctly
early on during the mission. Similarly, near- identify signals with amplitude close to the mea-
Earth-asteroid information will be communicated surement uncertainties, particularly in the pres-
at short notice. More information can be found in ence of larger signals induced by other
Lindegren (2010), while other organizational companions and/or sources of astrophysical
details and the latest news on payload and satel- noise of comparable magnitude. Finally, in
lite developments are available on the Gaia web cases of multiple-component systems where
pages at http://www.rssd.esa.int/gaia/. dynamical interactions are important (a situation
Gaia Mission 909
experienced already by radial-velocity surveys), revisited earlier findings using a more realistic
fully dynamical (Newtonian) fits involving an double-blind protocol. In this particular case, sev-
n-body code might have to be used to properly eral teams of “solvers” handled simulated
model the Gaia astrometric data and to ensure the datasets of stars with and without planets and
short- and long-term stability of the solution (see independently defined detection tests, with levels
Sozzetti 2005). of statistical significance of their choice, and
All the above issues could have a significant orbital fitting algorithms, using any local, global,
impact on Gaia’s capability to detect and charac- or hybrid solution method that they judged was
terize planetary systems. For these reasons, a best. The solvers were provided no information
Development Unit (DU) has been specifically on the actual presence of planets around a given
devoted to the modeling of the astrometric sig- target.
nals produced by planetary systems. The DU is A double-blind test campaign to estimate the
composed of several tasks, which implement potential of Gaia for characterizing planetary sys-
multiple robust procedures for (single and multi- tems (Casertano et al. 2008) showed that the fol- G
ple) astrometric orbit fitting (such as Markov lowing could be accurately modeled: (a) planets
Chain Monte Carlo and ▶ genetic algorithms) having an effect on the astrometric signature
and the determination of the degree of dynamical (a expressed in m as) with a value around six
stability of multiple-component systems. times the single-measurement error (s expressed
in m as) and orbital periods shorter than the nom-
inal 5-year mission lifetime and (b) favorable
Key Research Findings configurations of two-planet systems with well-
separated periods (both planets with a period
Gaia’s mode of operation (a signal-to-noise lim- inferior to 4 years, a ratio a/s > 10, and redun-
ited survey with uneven coverage, including time dancy over a factor of 2 in the number of observa-
sampling and scanning geometry, depending on tions). In the latter case, it is possible to carry out
ecliptic latitude) is such that there cannot be any meaningful coplanarity tests (relative inclination
optimization to the case of extrasolar planets. The of more than 10 ).
fundamental requirement, i.e., to have sufficient Overall, Casertano et al. concluded that Gaia
astrometric accuracy at magnitudes brighter than astrometry could allow the discovery and mea-
V = 13, was established at the time of the science surement of massive ▶ giant planets
case definition. Since little can be done with the (Mp > 2–3 MJ) with an orbit semimajor axis
photometric and spectroscopic capabilities between 1 and 4 astronomical units (au) orbiting
aboard the satellite, which cannot compete with solar-type stars as far as the nearest star-forming
present and planned ground-based facilities for regions. Saturn-mass planets with similar orbital
very-high-precision radial-velocity measure- semimajor axes around late-type stars within
ments (Pepe and Lovis 2008) and spaceborne 30–40 pc (see Fig. 1) could also be detected.
observatories devoted to ultrahigh precision tran- From these results, using Galaxy models and
sit photometry (e.g., Sozzetti et al. 2010), the with the current knowledge of frequency of
potential contribution of Gaia to exoplanet sci- exoplanets, it is possible to infer the number of
ence must be primarily gauged in terms of its planets of given mass and orbital separation that
astrometric capabilities. can be detected and characterized by Gaia.
A number of authors have tackled the problem Table 1 demonstrates that Gaia’s main strength
of evaluating the sensitivity of the astrometric will be its ability to accurately measure orbits and
technique required to detect extrasolar planets masses for thousands of giant planets and to per-
and reliably measure their orbital elements and form coplanarity measurements of a few hundred
masses (Sozzetti 2005, and references therein). multiple systems with favorable configurations.
The two most recent exercises on this subject Sozzetti et al. (2014) have recently revisited
(Casertano et al. 2008; Traub et al. 2010) have the topics of giant planet detection and
910 Gaia Mission
104
103
102
Planet mass (ME)
101
100
10−1
10−2
0.01 0.10 1.00 10.00
Semi-major axis (Au)
Gaia Mission, Fig. 1 Exoplanet discovery space for the assumptions are sV = 5 103 mag (upper curve)
Gaia astrometry, Doppler, and transit techniques. Detect- and sV = 1 105 mag (lower curve), S/N = 9, M* = 1
ability curves are defined on the basis of a 3s (standard MSUN, R* = 1 RSUN, uniform and dense (>1,000 data
deviation) criterion for signal detection. The blue curves points) sampling. Pink dots indicate the inventory of
are for Gaia astrometry with sA = 15 m as, assuming a Doppler-detected exoplanets as of December 2008.
1 MSUN G dwarf primary at 200 pc and a 0.4 MSUN Transiting systems are shown as light-blue filled dia-
M dwarf at 25 pc, respectively. The survey duration is monds, while the red hexagons are planets detected by
set to 5 years. The radial-velocity curves (red lines) microlensing. Solar System planets are also shown as
assume sRV = 3 m/s (upper curve) and sRV = 1 m/s green pentagons. The small yellow dots represent a theo-
(lower curve), M* = 1 MSUN, and a 10-year survey dura- retical distribution of masses and final orbital semimajor
tion. For visible-light transit photometry (green curves), axes from Ida and Lin (2008)
Gaia Mission, Table 1 Top: number of giant planets that could be detected (Nd) and measured by Gaia, as a function of
increasing distance (Dd). Star counts (Ns) are obtained using models of stellar population synthesis (Bienaymé
et al. 1987), while the Tabachnik and Tremaine (2002) model for estimating planet frequency as a function of mass
(DM p) and orbital period (Da) is used. Bottom: number of planetary systems that Gaia could potentially detect and
measure and for which coplanarity tests could be carried out successfully
Dd (pc) Ns Da (AU) DMp (MJ) Nd Nm
0–50 1 104 1.0–4.0 1.0–13.0 1,400 700
50–100 5 104 1.0–4.0 1.5–13.0 2,500 1,750
100–150 1 105 1.5–3.8 2.0–13.0 2,600 1,300
150–200 3 105 1.4–3.4 3.0–13.0 2,150 1,050
Case No. of systems
Detection 1,000
Orbits and masses (<15 % accuracy) 400–500
Coplanarity tests 150
Gaia Mission 911
characterization with Gaia focusing on the sam- content in the protoplanetary disk will be
ple of nearby low-mass M dwarf stars. They probed with unprecedented statistics, thanks
found that, given present-day estimates of the to the thousands of metal-poor stars and hun-
fraction fp of Jupiter-mass companions to dreds of young stars screened for giant planets
M dwarfs, comprehensive screening by Gaia of out to a few AU
the reservoir of M dwarfs within 100 pc could 3. Achieve key improvements in our comprehen-
result in ~2,600 new detections and as many as sion of important aspects of the formation and
500 accurate orbit determinations. It would then dynamical evolution of multiple-planet sys-
be possible to provide estimates of fp over ten tems: for example, the measurement of orbital
times more precise than those available to date. parameters for hundreds of multiple-planet
Sozzetti et al. (2014) also determined that Gaia systems, including meaningful coplanarity
astrometry, by precisely measuring the orbital tests, will allow discrimination between vari-
inclination, could alert us of the existence of ous proposed mechanisms for dynamical
tens of potentially transiting long-period planets. interaction G
Finally, they gauged the ability of Gaia to accu- 4. Aid in the understanding of direct detections
rately predict the ephemerides of (transiting and of giant extrasolar planets: for example, actual
non-transiting) planets around M stars and its mass estimates and full orbital geometry
potential to help in the precise determination of determination for suitable systems will inform
the emergent flux, for direct imaging and system- direct imaging surveys about the epoch and
atic spectroscopic characterization of their atmo- location of maximum brightness, in order to
spheres with dedicated observatories from the estimate optimal visibility, and will help in the
ground and in space. modeling and interpretation of giant planets’
phase functions and light curves
5. Provide important supplementary data for the
Applications optimization of the target selection for future
observatories aiming at the direct detection
Gaia’s main contribution to exoplanet science and spectral characterization of habitable ter-
will be its unbiased census of planetary systems restrial planets: for example, all F, G, K, M
orbiting hundreds of thousands nearby stars within the useful distance of 25 pc will
(d < 200 pc), relatively bright (V < 13) stars be screened for Jupiter- and Saturn-sized
across all spectral types, screened with constant planets out to several AU. These data will
astrometric sensitivity. The Gaia data have the help probe the long-term dynamical stability
potential to: of their habitable zones, where terrestrial
planets may have formed and maybe found
1. Significantly refine our understanding of the
statistical properties of extrasolar planets: the
predicted database of several thousand extra- Future Directions
solar planets with well-measured properties
will allow, e.g., to test the fine structure of An improvement of two to three orders of mag-
giant planet parameters’ distributions and fre- nitude in achievable measurement precision,
quencies and to investigate their possible down to the m as level, would allow this technique
changes as a function of stellar mass, to achieve in perspective the same successes as
metallicity, and age with unprecedented the Doppler method, for which the improvement
resolution from the km/s to the m/s precision opened the
2. Help crucially test theoretical models of gas doors for groundbreaking results in exoplanetary
giant planet formation and migration: for science. Indeed, m as astrometry is almost coming
example, specific predictions on formation of age. The largest compilation of astrometric
timescales and the role of varying metal orbits of giant planets (in many cases, signposts
912 Galactic Habitable Zone
of more interesting systems!), unbiased across all of giant planets around nearby M Dwarfs: the Gaia
spectral types up to d 200 pc, will allow Gaia potential. MNRAS 437:497–509
Tabachnik S, Tremaine S (2002) Maximum-likelihood
to crucially contribute to several aspects of plan- method for estimating the mass and period distribu-
etary system astrophysics (formation theories, tions of extrasolar planets. MNRAS 335:151–158
dynamical evolution), in combination with Traub WA et al (2010) Detectability of terrestrial planets
present-day and future extrasolar planet search in multi-planet systems: preliminary report. EAS Publ
Ser 42:191–199 (arXiv:0904.0822)
programs.
See Also
Galactic Habitable Zone
▶ Giant Planets Nikos Prantzos1 and Leticia Carigi2
▶ Hipparcos 1
▶ Lagrangian Points 2
Instituto de Astronomı́a, Universidad Nacional
▶ Parallax Autónoma de México, México, DF, Mexico
▶ Planetary Migration
▶ Proper Motion
Keywords
Habitable zone; Intelligent life; Extraterrestrial
Bienaymé O, Robin AC, Crézé M (1987) The mass den- civilization
sity in our galaxy. Astron Astrophys 180:94–110
Casertano S, Lattanzi MG, Sozzetti A et al (2008) Double-
blind test program for astrometric planet detection Definition
with Gaia. Astron Astrophys 482:699–729
ESA (1997) The Hipparcos and Tycho Catalogues, ESA
SP-1200 A galactic habitable zone is the putative region
ESA (2000) Gaia: composition, formation, and inside a galaxy with physical conditions compat-
evolution of the galaxy, Technical report. ESA-SCI ible with the origin, development, and long-term
(2000)4
Ida S, Lin DNC (2008) Toward a deterministic model of existence of life-as-we-know-it. Minimum
planetary formation. V. Accumulation near the ice line requirements are the existence of enough heavy
and super-earths. Astrophys J 685:584–595 chemical elements to form Earth-like planets and
Lindegren L (2010) Gaia: astrometric performance and a low occurrence of catastrophic events (e.g.,
current status of the project. Proc IAU Symp
261:296–305 supernovae, close stellar encounters) in order to
Pepe F, Lovis C (2008) From HARPS to CODEX: explor- allow the evolution of complex life forms.
ing the limits of Doppler measurements. Phys Scr
130:014007
Perryman MAC et al (2001) GAIA: composition, forma-
tion and evolution of the galaxy. Astron Astrophys Overview
369:339–363
Sozzetti A (2005) Astrometric methods and instrumenta- The idea underlying the concept of galactic
tion to identify and characterize extrasolar planets: a ▶ habitable zone (GHZ) is that various physical
review. PASP 117:1021–1048
Sozzetti A (2010) Detection and characterization of plan- processes, which may favor the development or
etary systems with mas astrometry. EAS Publ Ser the destruction of life, may depend strongly on
42:55–77 (arXiv:0902.2063) the temporal and spatial position in a galaxy. For
Sozzetti A et al (2010) Blue Dots team transits working instance, the risk of a ▶ supernova
group review. ASP Conf Ser 430:45–54
(arXiv:0912.0887) (SN) explosion, sufficiently close to represent a
Sozzetti A, Giacobbe P, Lattanzi MG, Micela G, threat for life, is in general larger in the inner
Morbidelli R, Tinetti G (2014) Astrometric detection galaxy than in the outer one and so is the
Galaxy 913
metallicity (for astronomers, the abundance of ▶ Habitable Zone

elements heavier than helium) of the interstellar ▶ Milky Way
medium, which may be important for the forma- ▶ Supernova
tion of Earth-like exoplanets. Assuming that all
relevant factors (hostile or beneficial to complex
life) can be properly quantified, one may use
models of galactic evolution in order to evaluate Carigi L, Garcı́a-Rojas J, Meneses-Goytia S (2013) Chem-
the position and extent of the GHZ over time. ical evolution and the galactic habitable zone of M31,
Preliminary attempts gave, up to now, controver- the Andromeda Galaxy. Rev Mex Astron Astrofis
sial results as to the extent of the GHZ of the 49:253–273
Gowanlock MG, Patton DR, McConnell SM
▶ Milky Way disk, because of the difficulty in (2011) A model of habitability within the Milky Way
properly evaluating the relevant factors, mainly galaxy. Astrobiology 11:855–873
the chemical requirement to form terrestrial Jiménez-Torres JJ, Pichardo B, Lake G, Segura A (2013)
planets. It is hard to quantify the role of Habitability in different Milky Way stellar environ- G
ments: a stellar interaction dynamical approach. Astro-
metallicity in planetary formation and even biology 13(5):491–509
harder to draw any quantitative conclusions Lineweaver CH, Fenner Y, Gibson BK (2004) The galac-
about the probability of definitive sterilization of tic habitable zone and the age distribution complex life
a habitable planet. Life displays unexpected in the Milky Way. Science 303:59–62
Prantzos N (2011) The concept of “galactic habitable
robustness and a cosmic catastrophe might even zone”. In: Gargaud M, Lopez-Garcı́a P, Martı́n
accelerate evolution toward life forms that are H (eds) Origins and evolution of life: an
presently unknown. Studies of GHZ focused astrobiological perspective. Cambridge University
mainly on the Milky Way disk, while the Milky Press, Cambridge, pp 154–166
Sundin M (2006) The galactic habitable zone in barred
Way halo (presumably too metal poor to allow galaxies. Int J Astrobiol 5:325–326
for planetary formation) and bulge (presumably Suthar F, McKay CP (2012) The galactic habitable zone in
too dense to avoid frequent catastrophes there) elliptical galaxies. Int J Astrobiol 11:157–161
appear – perhaps incorrectly – as less interesting
abodes for complex life. The GHZ has recently
been computed for two elliptical galaxies and the
disk of Andromeda. Very few of those studies Galaxy
include cinematic and dynamic aspects of the
stars and their planetary systems. These issues Nikos Prantzos
are fundamental to infer galactic regions where Institut d’Astrophysique de Paris, Paris, France
potentially habitable planets form, if stars have
migrated. In addition, the stability of planetary
orbits can be affected by the proximity of stars. Definition
The concept of GHZ is much more poorly defined
than the one of circumstellar habitable zone. At A galaxy is a massive, gravitationally bound sys-
present, it should be considered, at best, as a tem of ▶ stars, which may also include stellar
broad framework, allowing us to formulate our remnants (white dwarfs, neutron stars, black
thoughts about a very complex phenomenon such holes), gas, and dust, as well as nonbaryonic
as life (origin, development, and survival) in a dark matter. The morphological classification of
galactic environment. galaxies, established by E. Hubble in the 1930s,
includes ellipticals, spirals, and irregulars, each
class divided in subclasses and going from “early
See Also types” (generally more massive and gas poor,
which is the case for ellipticals) to “late types”
▶ Abundances of Elements (less massive and gas rich, as the irregular galax-
▶ Exoplanets, Discovery ies). Galaxies are generally found in groups (up to
914 Galilean Satellites
a few tens of members), clusters (a few thou- nights. Those four biggest Jupiter’s moons, or
sands), and superclusters. Our own ▶ Milky Medicean Stars as Galileo named them, are: Io,
Way, often called the Galaxy, is a large spiral Europa, Ganymede, and Callisto (today Jupiter
with a baryonic mass of 51010 MJ, slightly registers 50 moons and 16 provisional moons).
smaller than the largest of the Local Group gal- The Galilean satellites are quite different from
axies (M31 or Andromeda). There is no univer- each other. Ganymede is the largest moon in our
sally accepted definition of a galaxy, nor a clear Solar system and is the only moon known to have
lower limit on its stellar mass: some systems its own internally generated magnetic field. Io has
characterized as “galaxies” (tidal dwarfs, a volcanic nature. The surface of Europa is
ultracompact dwarf spheroidals) are less massive mostly water ice (it is believed that it has twice
than the most massive ▶ globular clusters. the water existing on Earth), and Callisto shows
an extremely heavily cratered and ancient
surface.
See Also
▶ Globular Cluster
▶ Milky Way
Galileo Galilei
▶ Star
Therese Encrenaz
Galilean Satellites
Fernando B. Figueiredo Definition

Portugal Galileo Galilei (1564–1642) is one of the
most famous astronomers over the whole
history of humanity. With the telescope he
Synonyms designed, he was able to make major discoveries
that supported and reinforced the new
Jupiter’s biggest moons; Medicean Stars heliocentric theory proposed by Nicolaus Coper-
nicus that drastically modified our view of the
world.
Definition
At the night of 7 January 1610 the Italian physi- Overview

cist and mathematician Galileo Galilei
(1564–1642) observed the planet Jupiter with a Galileo, born in Pisa in Italy, started to learn
telescope made by him and saw what he thought medicine and then turned to mathematics. After
were three fixed stars near it. In subsequent obser- a first stay in Pisa, in 1592 he got a chair of
vations made in the following nights Galileo mathematics at Padua University. In 1604, on
found that the bright spots were not three but the occasion of the apparition of a new star in
four and not fixed stars but rather planetary bod- Sagittarius, he gave three public lectures on the
ies that revolved around Jupiter, since their posi- “Nova,” showing from the absence of ▶ parallax
tions related to the planet had changed over those that the star was at a great distance from Earth and
Galileo Mission 915
thus challenging Aristotle’s views about the solar system. By computing the exact periods and
unchangeability of celestial bodies. the times of mutual ▶ occultation of the four
In 1609, Galileo took advantage of new satellites, Galileo provided navigators with a
research about spyglasses in Holland to develop method for measuring the longitude from any-
the first concept of an astronomical telescope. In where at sea. Later, the exact timing of Io’s
August 1609, he presented the Doge of Venice occultations allowed the astronomer Olaf
with his first instrument. In January 1610, with a Roemer to get a first estimate of the speed of
20-times magnifying instrument, he made the light; his measured speed was 30 % lower than
historical discovery of the four big satellites of the presently accepted value.
▶ Jupiter, which he called “Medicean” in honor
of Cosimo de Medici, Duke of Tuscany (they are
now known as the Galilean satellites). This dis- See Also
covery, together with the irregular shape of the
Moon’s surface and the multitude of stars ▶ Callisto G
forming the Milky Way, was reported in the ▶ Europa
historical manuscript “Sidereus Nuncius,” ▶ Galilean Satellites
published early in 1610. Still in 1610, Galileo ▶ Ganymede
observed Venus’ phases, supporting a heliocen- ▶ Io
tric solar system, moving sunspots, and the ▶ Jupiter
changing shape of Saturn that he was not able to
interpret, but which was in fact due to Saturn’s
rings. These results were reported in Galileo’s
Discourse on Bodies in Water in 1612 and “Letter Mc Mullin E (2001) Galileo Galilei. In: Murdin P (ed) The
on Sunspots” in 1613. encyclopedia of astronomy and astrophysics. IoP,
In 1616, after the Roman Congregation of the Bristol, pp 921–925
Index banned Copernicus’s De Revolutionibus as Swerdlow NM (1998) Galileo’s discoveries with the tele-
scope and their evidence for the Copernican theory. In:
being “contrary to Scriptures,” Galileo was Machamer P (ed) The Cambridge companion to Gali-
forced to abandon his plans for a Copernican leo. Cambridge University Press, Cambridge,
treatise. He came back to resume his work in pp 244–270
1623, when Pope Urban VIII, an admirer of
Galileo’s work, became elected. However, in
1632, his Dialogue Concerning the Two Chief
Worlds Systems raised the hostility of the Galileo Mission
Roman theologians; Galileo was summoned to
Rome and put on trial. He was forced to abjure Michel Viso
publicly the prohibited claims of the Earth’s CNES/DSP/SME, Vétérinaire/DVM,
motion and the Sun’s being at rest and was sen- Astro/Exobiology, Paris, Cedex 1, France
tenced to house arrest. His last manuscript “Two
New Sciences,” published in Holland in 1638,
was devoted to mechanics. Definition
Among the many discoveries that were made
possible by Galileo’s telescope, the discovery of The ▶ NASA Galileo mission was launched from
the four Galilean satellites probably had the most Cape Canaveral on October 18, 1989, on board
important implications for astronomy. In contrast the Space Shuttle Atlantis. The spacecraft and the
with Aristotle’s doctrine, they demonstrated that inertial upper stage were released and fired for a
there is more than one center of rotation in the 6-year trip in the solar system, heading to Venus,
916 Galileo Mission
then to the Earth to perform three gravity-assisted order to avoid any possible crash on, and contam-
maneuvers, before finally cruising toward ination of, Europa. Galileo entered the Jovian
▶ Jupiter. atmosphere September 21, 2003, at 19 h
49 GMT after 35 orbits. Europa and its ocean
constitute one of the major targets of relevance
Overview for astrobiology (Figs. 1 and 2).
On December 7, 1995, a Jupiter probe released

from Galileo entered the Jovian atmosphere with
a parachute and transmitted data, relayed by the
main spacecraft, for more than 59 min. Then,
using the main engine, Galileo was inserted in
an elliptic 2-month-long orbit around Jupiter.
During the primary (11 orbits) and the extended
(14 orbits) mission, Galileo flew by and analyzed
most of the Jovian satellites. The mission was
extended several more years. Galileo made
numerous discoveries about Jupiter and its
moons. One of the most striking was the evidence
of an ocean below the icy crust of ▶ Europa. The
measures of the induced ▶ magnetic field for
Europa (as for ▶ Callisto) while orbiting through
the strong Jovian magnetic field were consistent
with the presence of conducting layers. This fea-
ture may best be explained by the presence of
deep salty liquid-water oceans, for which there
was already indirect geological evidence (in the
case of Europa, see references). Because of the
potential existence of such an ocean, NASA
decided to destroy the spacecraft at the end of Galileo Mission, Fig. 1 The Galileo spacecraft prepara-
the mission by entering Jupiter’s atmosphere, in tion at Kennedy Space Centre (Photo NASA)
Galileo Mission,
Fig. 2 Europa mosaic
image in true and false
colors (Photo Nasa)
Gamma Cephei 917
Reference and Further Reading and, at the same time, very surprising discovery.
It was exciting because if true, it would have
Khurana KK, Kivelson MG, Stevenson DJ, Schubert G, marked the detection of the first planet outside
Russell CT, Walker RJ, Polanskey C (1998) Induced
of our solar system. It was surprising because the
magnetic fields as evidence for subsurface; oceans in
Europa and Callisto. Nature 395:777–780 planet-hosting star is the primary of a binary
system with a separation of ~20 AU, a distance
comparable to the planetary distances in our solar
system.
Gamma Cephei The moderately close orbit of the stellar com-
panion of g Cephei raised questions about the
Nader Haghighipour reality of its planet. The skepticism over the
Institute for Astronomy, University of Hawaii- interpretation of the results (which was primarily
Manoa, Honolulu, Hawaii, HI, USA based on the idea that binary star systems with
small separations would not be favorable places G
for planet formation) became so strong that in a
Definition subsequent paper in 1992, Walker and his col-
leagues suggested that the planet in the g Cephei
Gamma Cephei (g Cephei, g Cep) is a binary star binary might not be real, and the variations in the
system in which the primary star, Gamma radial velocity of this star might have been due to
Cephei A, is orbited by a Jupiter-mass planet. its chromospheric activities.
Despite the 1992 article, the search for planets
in binaries did not stop. Gamma Cephei was
Overview continuously monitored and more precise mea-
surements of its radial velocity variations were
In 1988, in an article on the analysis of the mea- obtained. In 2003, 15 years after the first
surements of the variations in the radial velocities announcement of the planet of this system, these
of a number of stars, Campbell, Walker, and efforts bore fruit, and in an article in Astrophys-
Yang reported an interesting phenomenon; the ical Journal, Hatzes and his colleagues con-
radial velocity variations of g Cephei seemed to firmed the existence of a Jupiter-like planet
suggest the existence of a Jupiter-like planet around the primary of g Cephei (Fig. 2). As the
around this star (Fig. 1). This was a very exciting planet became real, it introduced many
Gamma Cephei,
Fig. 1 Radial velocity 100
residuals for g Cephei after g Cep Residuals
the subtraction of the
second order fit to the
Velocity (m s−1)
original radial velocity data

(Campbell et al. 1988)
−100
1981 1982 1983 1984 1985 1986 1987
Year
918 Gamma Cephei
1985 1990 1995 2000

100
80 γ Cephei CFHT
McD I
McD II
Relative velocity (meters/second)
McD III
60
40
20
−20
−40
−60
45000 46000 47000 48000 49000 50000 51000 52000 53000

JD - 2400000 (days)
γ Cephei
100
80 CFHT McD I
60
40
20
Relative velocity (meters/second)
0
−20
−40
−60
100
80 McD II McD III
60
40
20
0
−20
−40
−60
0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0
Orbital Phase
Gamma Cephei, Fig. 2 Radial velocity measurements of g Cephei as reported by Hatzes et al. (2003) obtained at the
CFHT (Canadian, French, Hawaii Telescope) and the McDonald observatory
Gamma Rays 919
Haghighipour N (2010) Planets in binary star systems,

vol 366, Astrophysics and space science library.
Springer, Dordrecht
Hatzes AP, Cochran WD, Endl M, McArthur B, Paulson
DB, Walker GAH, Campbell B, Yang S (2003)
A planetary companion to gamma Cephei
A. Astrophys J 599:1383–1394
Gamma Rays
Kazumichi Nakagawa
Graduate School of Human Development and
Environment, Kobe University, Nada, Kobe, G
Japan
Gamma Cephei, Fig. 3 Schematic view of the binary

star and planetary orbits of g Cephei system
Keywords
challenges to planetary science. Figure 3 shows a Electromagnetic wave; Radiation

schematic figure of this system.
The 2003 confirmation of g Cephei’s planet,
and the subsequent detection of giant planets in Synonyms
other moderately close binary stars such as GL
86, HD 41004, HD 196885, HD 176051, and a High-energy photons; High-energy radiation
Centauri, marked the beginning of a new era in
theoretical and observational research on planets
in dual-star systems. During the past few years, Definition
much research has been carried out in this area,
and a large number of excellent articles have been Gamma rays are electromagnetic waves with
published on different aspects of observational high photon energy. Gamma rays are emitted
and theoretical studies of planets in moderately (1) via transitions between quantum levels of
close binaries. For more details, see the book, nuclei and elementary particles, (2) via
Planets in Binary Star Systems (Haghighipour particle–antiparticle annihilation and (3) via
2010). bremsstrahlung (photon emission due to interac-
tion between the strong magnetic field of nuclei
and high-energy particles in the vicinity of the
See Also nuclei). Gamma rays interact with matter through
(1) photoelectron emission, (2) Compton scatter-
▶ Binary Stars, Young ing with electrons, (3) creation of
▶ Exoplanets, Discovery electron–positron pairs, and (4) nuclear
▶ Radial-Velocity Planets photoreactions.
References and Further Reading Overview

Campbell B, Walker GAH, Yang S (1988) A search for
substellar companions to solar-type stars. Astrophys As the result of the interaction of a gamma ray
J 331:902–921 with matter, the gamma ray photon is annihilated
920 Ganymede
and its energy is transferred to an electron. The See Also

resulting high-energy electron will interact with
other electrons in the matter to ionize or excite ▶ Radiation Biology
molecules or atoms and will gradually dissipate ▶ Radiochemistry
its energy to produce large numbers of ▶ Radiolysis
high-energy electrons in the system. Subsequent
radiation chemistry due to these high-energy
electrons will be responsible for chemical pro-
cesses. Chemical evolution or molecular damage Bonner WA, Hall H, Chow G, Liang Y, Lemmon RM
are the result of this radiation chemistry. (1985) The radiolysis and radioracemization of amino
According to Platzman (1962), the magnitude of acids on clays. Orig Life 15:103–114
the radiation chemical effect G due to an electron Kaneko F, Tanaka M, Narita S, Kitada T, Matsui T,
Nakagawa K, Agui A, Fujii K, Yokoya A (2005)
with initial kinetic energy E0 is expressed as Chemical evolution of amino acid induced by soft
ð E0 X-ray with synchrotron radiation. J Electron Spectrosc
1 df Relat Phenom 144–147:291–294
G/ dE Platzman RL (1962) Superexcited states of molecules, and
IP E dE
the primary action of ionizing radiation. Vortex
23:372–385
where IP is the ionization potential of the system
and df/dE the optical oscillator strength distribu-
tion, which is proportional to the absorption cross
sections.
In order to evaluate the radiation chemical Ganymede
effect caused by gamma rays, the energy effi-
ciency G-value (number of chemical events per Therese Encrenaz
100 eV absorbed) is essential, instead of the pho- LESIA, Observatoire de Paris - Section de
ton efficiency or quantum yield (number of chem- Meudon, Meudon, France
ical events per one absorbed photon). For
radiation chemistry, the magnitude of the
G-value should be proportional to the incident Keywords
photon energy hn of the gamma rays because
the incident photon energy is divided into the Galilean satellites
small value of IP. The value of hn/IP is about
105 in the case of hn = 1 MeV and IP = 10 eV.
On the contrary, for photochemistry in which Definition
photon energy is ordinarily smaller than 10 eV,
the reaction cannot occur if the photon energy is Ganymede is the third Galilean satellite in terms
not resonant with the excited state which is of its distance from Jupiter; on the other hand, it is
responsible for the characteristic transition. the biggest satellite in the solar system: it is
Chemical effects induced by higher-energy bigger than the planet Mercury. Like the other
photons such as gamma rays, X-rays, and ultra- Galilean satellites, Ganymede was discovered in
violet light are similar to each other. It is known January 1610 when Galileo Galilei first pointed
that the chemical effects include decomposition, his refractor toward Jupiter. It was named by
dimerization, polymerization, and ionization. Simon Marius (1614) after a lover of Zeus, who
Gamma ray-induced racemization and decompo- corresponds to Jupiter in Greek mythology. Gan-
sition of amino acids was reported by Bonner ymede’s average distance to Jupiter is slightly
et al. (1985). Dimer formation of alanine mole- above 1 million kilometres or about 15 Jovian
cules induced by X-rays was reported by Kaneko radii. Ganymede’s density is 1.9 g/cm3, typical
et al. (2005). of a 1:1 mixture of water ice and rocks. Water ice
Gas Chromatography 921
was identified at Ganymede’s surface as early as and possibly from tidal heating (although less
1971 through ground-based near-infrared efficiently than in the case of ▶ Io and ▶ Europa)
spectroscopy. in Ganymede’s past history.
In 2012, ESA selected a space mission
devoted to the exploration of Ganymede and the
Overview Galilean satellites, with the main purpose of
exploring habitability conditions in icy moons.
The space exploration of Ganymede started with The JUICE mission (JUpiter and ICy moons
the Voyager 2 flybys in 1979 and was followed Explorer) is planned for a launch in 2022. It will
after 1995 by the Galileo orbiter that operated approach Jupiter in 2030 and, after a series of
until 2003. flybys, will enter the orbit around Ganymede in
The surface of Ganymede, as revealed by 2033 for an in-depth monitoring of the satellite.
Voyager and Galileo images, is composed of
dark and bright terrains. The old dark terrains G
are heavily cratered and made up of patchy fea- See Also
tures of different albedos; the dark material prob-
ably corresponds to organic deposits provided by ▶ Cryovolcanism
asteroid and comet impacts. The dark regions are ▶ Europa
fractured by tectonic activity and impact craters, ▶ Io
many of them being the signature of the ▶ late ▶ Jupiter
heavy bombardment. Bright terrains show
smooth, young areas with grooves and stripes
resulting from tectonic activity. These terrains
cover three quarters of Ganymede’s surface and Schubert G, Klemaszewsky J (2001) Ganymede. In:
seem to have been resurfaced mostly by tectonic Murdin P (ed) The encyclopedia of astronomy and
activity and by active (▶ cryovolcanism). astrophysics. IoP, Bristol
The unexpected discovery of an intrinsic mag-
netic field on Ganymede, by the radio science
instrument and the magnetometer of the Galileo
mission, strongly suggests that the satellite’s inte- Gas Chromatography
rior is differentiated and contains a melted iron
nucleus at the center. This magnetic field induces Jennifer C. Stern
a small stable ▶ magnetosphere 5,000 km in Planetary Environments Laboratory, NASA
radius where high-energy particles, radiation Goddard Space Flight Center, Greenbelt,
belts, and auroras have been observed. Gany- MD, USA
mede’s magnetosphere, generated by its magnetic
field, is in electrodynamical interaction with the
Jovian magnetic field through a permanent mag- Synonyms
netic reconnection. It acts like a shield that allows
the existence of an extended ionosphere, GC
uncompressed by the Jovian magnetic flux.
The interior of Ganymede appears to be dif-
ferentiated in three layers, with a central metallic Definition
core, a silicate mantle, and a water-ice-rich man-
tle and crust. Convection through the icy outer Gas chromatography is a laboratory analytical
layer transfers the internal heat to the surface. The method for separating volatile components of a
internal heat comes from radioactive elements mixture as they are passed in a gaseous mobile
within the silicate mantle, from gradual cooling, phase past a stationary phase or through a
922 Gas Chromatography/Mass Spectrometry
column. Separation of molecules is based on dif- dynamical interactions with the surrounding gas-
ferences in the affinity of species for the mobile eous material act in the opposite direction to the
and stationary phases, which may be influenced motion of the body. Gas drag is primarily impor-
by polarity, molecular weight, and volatility. Gas tant in planetary dynamics because it functions as
chromatography is often coupled with a detector one of the main processes that remove energy
to quantify and/or identify molecular species. from orbiting bodies in a dense gaseous ▶ proto-
Detection methods include flame ionization, ther- planetary disk, where it causes loss of orbital
mal conductivity, and mass spectrometry. Gas momentum and decrease in orbital eccentricity,
chromatography has been used for organic anal- inclination, and semimajor axis of a body in
ysis on Mars missions, including ▶ Viking and ▶ orbit around the central star. Gas drag is more
the 2011 Mars Science Laboratory (MSL). significant for small (less than 100 km) bodies.
See Also
See Also
▶ Orbit
▶ Chromatographic Coelution ▶ Protoplanetary Disk
▶ Chromatography
▶ GC/MS
▶ MSL
▶ Viking
Gas Giant Planet
Daniel Rouan
Gas Chromatography/Mass Meudon, France
Spectrometry
▶ GC/MS Definition
A gas giant planet is an expression often used to

designate the class of extrasolar planets “that are
Gas Drag of a type analog” Jupiter and Saturn, to the most
massive giant planet in the solar system. They are
Avi M. Mandell characterized by a big mass (say larger than
NASA Goddard Space Flight Center, Greenbelt, 100 times the Earth’s mass) which implies that
MD, USA they must have an extended gaseous envelope
and a rather low density. Indeed, in the various
cases where both the mass and the size were
Keywords measured, massive exoplanets feature a density
well within the expected range, i.e., one of the
Migration solar system’s giant planets (0.7–1.7 g cm3).
Definition See Also
Gas drag is a force operating on a body moving ▶ Giant Planets

through a gaseous medium, which causes the ▶ Jupiter
body to lose momentum and energy. The ▶ Saturn
Gaspra 923
more complex molecules by gas-phase ▶ neutral-

Gas-Grain Chemistry neutral and ▶ ion-neutral reactions.
Steven B. Charnley
Solar System Exploration Division, Code See Also
Space Flight Center, Greenbelt, MD, USA ▶ Interstellar Chemical Processes
Keywords ▶ Molecular Depletion
▶ Molecular Desorption
Molecular clouds; Interstellar chemical processes ▶ Molecules in Space
▶ Protostars
▶ Radical G
Definition
Gas-grain chemistry is the study of the
chemical processes involving solid dust grains Charnley SB, Rodgers SD (2009) Theoretical models of
embedded in interstellar/circumstellar gas. Gen- complex molecule formation on dust. In: Meech KJ,
erally, these are loss of atoms and molecules from Keane JV, Mumma MJ, Siefert JL, Werthimer DJ (eds)
the gas to the grains, catalytic reactions on sur- Bioastronomy 2007: molecules, microbes, and extra-
terrestrial life. Astron Soc Pacific, San Francisco,
faces, and return of surface species to the gas pp 29–34
phase. Tielens AGGM, Allamandola LJ (1987) Composition,
structure, and chemistry of interstellar dust. In:
Hollenbach DJ, Thronson HA (eds) Interstellar pro-
cesses. Reidel, Dordrecht, pp 397–470
Overview
There are four basic processes: (1) Gaseous atoms

and molecules collide with, and stick to, dust grains
at low temperatures. The net effect is the depletion Gaspra
of these species in the gas. (2) Adsorbed surface
species can migrate, meet, and react leading to new Stefano Mottola
▶ radical and closed shell molecules. Such reac- German Aerospace Center (DLR), Institute of
tions lead to the conversion of hydrogen atoms to Planetary Research, Berlin, Germany
molecules and to the growth of ice mantles on the
grain cores. These ices are dominated by water and
also contain CO, CO2, methanol, methane, and Definition
ammonia as well as more complex organic mole-
cules. (3) The third is desorption and the conse- (951) Gaspra is a main-belt asteroid named after a
quent return of surface species to the gas. Several Crimean town located on the Black Sea. It was
processes can remove material from the icy man- the first asteroid to be visited by spacecraft, when
tles, including photoejection by UV photons, ther- the NASA Galileo probe approached it on its
mal evaporation due to grain heating by cosmic ray course to Jupiter. The analysis of the Galileo
impacts or by photon absorption near ▶ protostars, images revealed a body approximately 18
and ▶ sputtering in ▶ shock waves. (4) The last is 10 9 km in size, with an irregular shape and
the chemistry in the hot gas into which whole or with a comparatively young surface age between
partial removal of the ice mantles has occurred. In 30,000 and 300,000 years. The body probably
this case, it is possible to synthesize many even originated in a disruptive collision that created
924 GC
the Flora asteroid family, which is believed to the chemical properties between the molecules in
comprise about 103 objects. Gaspra is classified a mixture cause them to migrate with different
as an S-type asteroid, and its surface displays the velocities as they travel the length of the column.
spectral signature of mafic minerals. Galileo As the molecules exit the GC, they are fed into
images confirmed the presence of subtle color the mass spectrometer where they are ionized and
variations across its surface, which had been pre- detected using their mass to charge ratio.
viously detected by ground-based observations These two components, used together, allow a
and which are interpreted as the manifestation much more precise identification of compounds
of downslope movement of regolith. than would be accomplished using either instru-
ment alone. Sometimes two different molecules
can have similar retention times, as measured by
GC, or have a similar mass spectrum, but it is
GC extremely unlikely that two different molecules
will have the same GC and MS response. Often a
▶ Gas Chromatography third dimension is added to GC/MS analysis,
either an extra GC column in GC/GC/MS or an
extra MS dimension in GC/MS/MS, both of
which further refine the quality of analysis.
GC/MS Several GC/MS instruments have been sent on
space missions. The Viking landers both carried
Henderson James (Jim) Cleaves II GC/MS instruments to Mars and the Huygens
Earth–Life Science Institute (ELSI), Tokyo probe studied the atmosphere of Saturn’s moon
Institute of Technology, Meguro–ku, Tokyo, ▶ Titan using GC/MS. However, the American
Japan Pioneer and Russian Venera 11 and 12 landers
Institute for Advanced Study, Princeton, analyzed Venus’ atmosphere using GC and MS
NJ, USA separately.
Washington, DC, USA
Center for Chemical Evolution, Georgia Institute See Also
▶ Huygens Probe
Synonyms ▶ Mass Spectrometry
Gas chromatography/mass spectrometry ▶ Titan
▶ Viking
Definition
In chemistry, gas chromatography/mass spec- GCM

trometry (GC/MS) is an analytical method that
combines a gas chromatograph and a mass spec- Lisa Kaltenegger
trometer. GC/MS was first developed in the Cornell University, Ithaca, NY, USA
1950s by Gohlke and McLafferty. The gas chro-
matography uses a capillary column, which may
have variable column dimensions (length and Synonyms
diameter) and stationary phase attributes
(composition and thickness). The differences in General circulation model; Global climate model
GEMs 925
Definition the horizon at this time). The gegenschein is

actually a slight brightness peak in the ▶ zodiacal
A general circulation model (GCM) is a complex light and is produced by back-scattering of sun-
mathematical model of the general circulation of light from ▶ interplanetary dust particles. The
a planetary atmosphere and/or ocean. A GCM is gegenschein is only visible under very dark sky
based on the Navier-Stokes equations on a rotat- conditions.
ing sphere with thermodynamic terms for various
energy sources (like radiation and latent heat).
GCMs are used for weather forecasting, under-
standing and projecting climate changes on
See Also
Earth, and, in simplified form, in exoplanet
▶ Exozodiacal Light
modeling.
▶ Interplanetary Dust Particle
▶ Zodiacal Light G
See Also
▶ Atmosphere, Model 1D GEMs

▶ Clouds William M. Irvine
▶ Exoplanets, Modeling Giant Planets’ University of Massachusetts, Amherst, MA, USA
Atmospheres
▶ Grey Gas Model
▶ Hadley Cells Definition
▶ Non-Grey Gas Model: Real Gas Atmospheres
The matrices of some interplanetary dust parti-
cles, particularly the anhydrous chondritic porous
(CP) class of ▶ interplanetary dust particles
(IDP), include glassy, submicron-sized silicate
grains that have been called GEMs (glasses
Gegenschein with embedded metal and sulfides). It has been
proposed that the GEMs are interstellar in origin
William M. Irvine and were incorporated into the IDPs or their par-
University of Massachusetts, Amherst, MA, USA ent bodies during the formation of the solar
system.
Synonyms
See Also
Counterglow
Definition
The gegenschein or counterglow is a faint

brightness enhancement in the night sky in the Bradley JP (1994) Chemically anomalous, preaccre-
direction opposite to that of the Sun (i.e., in the tionally irradiated grains in interplanetary dust from
antisolar direction; the Sun, of course, is below comets. Science 265:925–929
926 Gene
G.J. Mendel in his classic experiments aimed at

Gene studying the segregation of heritable traits in peas
and other garden plants. Later on, the Mendel’s
Carlos Briones factors were called “genes” and assigned to par-
Centro de Astrobiologı́a (CSIC/INTA), Consejo ticular positions or “loci” on the chromosomes
Superior de Investigaciones Cientı́ficas, Madrid, and DNA molecules. The different versions of
Spain the same gene are called alleles and, in diploid
species – those showing two copies of their
genetic material – one allele is inherited form
Keywords each parent. Diploid organisms can be homozy-
gous for a given gene if they contain two copies
Central dogma of molecular biology; Exon; Gene of the same allele or heterozygous if they possess
expression; Genome; Intron; mRNA; rRNA; a copy of two different alleles (Watson
Splicing; tRNA et al. 2008; Lewin 2008).
Genes are specific functional units of a
genome potentially transcribed into RNA which
Synonyms possibly will code for a protein. In the decade of
the 1940s, it was clear that specific genes code for
Cistron; Transcription unit specific proteins, thus leading to the so-called
“one gene-one enzyme” hypothesis. The determi-
nation of the ▶ genetic code permitted molecular
Definition biologists to unveil the correspondence between
coding nucleotide triplets – called
Various definitions of the term “gene” occurred “▶ codons” – in either the DNA gene or the
and still coexist. Genes are the units of heredity, transcribed mRNA and the amino acid they spec-
and this remains true since the pioneering contri- ify. It was further demonstrated that the majority
bution of G.J. Mendel in 1866. ▶ DNA is the of eukaryotic genes – and a low number of the
substrate of heredity, as it was shown by bacterial and archaeal ones – are not collinear
O.T. Avery and his collaborators in the 1940s; with their protein products. Instead, they are
thus, ▶ genetics and molecular biology have been split into segments called “▶ exons” that code
able to localize genes on this molecular substrate. for different protein fragments. Exons are sepa-
Nevertheless, some hereditary information lies rated by intervening sequences called “▶ introns”
also outside the classical or “true” genes. In (Roy and Gilbert 2006). After the ▶ transcription
cells, the classical genes are segments of their of the gene, a reaction called “RNA
DNA genome that can be transcribed into a mes- splicing” – either protein-catalyzed or
senger ▶ RNA, which will be finally translated autocatalyzed by the intron – allows for the exci-
into a protein, a ribosomal RNA, a transfer RNA, sion of the introns and the covalent joining of the
or other kinds of RNAs. Viral genes are frag- exons in a mature RNA molecule. If such a
ments of their DNA or RNA ▶ genome that, in mature RNA is an mRNA, further posttranscrip-
most cases, code for the structural or regulatory tional modifications are needed – including cap-
proteins that will be expressed by the molecular ping and polyA addition – before becoming ready
machinery of the infected cell. to be translated into the coded protein. Due to
their mosaic structure, discontinuous genes may
code for more than one protein if alternative
Overview splicing of their pre-mRNA occurs.
In addition to the transcribed sequences –
Inheritance in organisms occurs by means of fac- called “transcription units” or “units of
tors or “determinants,” as first concluded by transcription” – all genes have additional
Gene Expression 927
sequences, among them regulatory regions that ▶ Genome

control their expression. Thus, a common struc- ▶ Genotype
tural organization of a protein-coding eukaryotic ▶ Intron
gene can be summarized as follows: 50 -enhancer- ▶ Replication (Genetics)
promoter-exon1-intron1-exon2-. . .-exonN-termi- ▶ RNA
nation signal-polyadenilation sequence-down- ▶ Transcription
stream regulators-30 . In bacteria and archaea, ▶ Translation
introns are rare and the average length of genes ▶ Virus
is shorter. Two extreme examples of gene size are
the 21 nt-long gene mccA, coding for the
heptapeptide antibiotic microcin C7 in
Enterobacteria, and the 2.3 106 nt-long Hou Y, Lin S (2009) Distinct gene number-genome size
human dystrophin gene, which includes relationships for eukaryotes and non-eukaryotes: gene
79 exons and very long introns. The first content estimation for dinoflagellate genomes. PLoS G
sequenced gene was that of the bacteriophage ONE 4:e6978
Lewin B (2008) Genes IX. Jones and Bartlett, Boston
MS2 coat protein (Min Jou et al. 1972), an RNA Min Jou W, Haegeman G, Ysebaert M, Fiers W (1972)
▶ virus whose genome was also the first to be Nucleotide sequence of the gene coding for the bacte-
sequenced in its entirety. riophage MS2 coat protein. Nature 237:82–88
The number of genes in different genomes Roy SW, Gilbert W (2006) The evolution of spliceosomal
introns: patterns, puzzles and progress. Nat Rev Genet
ranges from 1 in the simplest viruses – viroids 7:211–221
lack genes in their genomes – to up to 105 in Snustad DP, Simmons MJ (2010) Principles of genetics,
certain eukaryotes. The relationship between 5th edn. Wiley, New Jersey
genome size and gene content has been investi- Watson J, Baker T, Bell S, Gann A, Levine M, Losick
R (2008) Molecular biology of the gene. Benjamin
gated based on the data from completely Cummings, Menlo Park
sequenced and annotated genomes. Viral, bacte-
rial, and archaeal genomes are mostly composed
of gene-coding sequences, with median values of
gene-coding percentages between 85 % and 90 %
in the three groups (range: 47–97 %). In turn, in Gene Expression
eukaryotes only a small proportion of their
genomes is ever transcribed as bona fide infor- Carlos Briones
mational RNAs – mRNA, rRNA, or tRNA “true” Centro de Astrobiologı́a (CSIC/INTA), Consejo
genes – showing a median gene-coding percent- Superior de Investigaciones Cientı́ficas, Madrid,
age of about 32 % (range: 1–82 %). For example, Spain
the gene content of the bacterium Escherichia
coli genome is 88 %, whereas in Homo sapiens
the coding sequences represent less than 2 % of Definition
our genome (Hou and Lin 2009; Snustad and
Simmons 2010). Gene expression is the overall process by which
information stored in a gene is used to produce a
gene product that may be either a protein, includ-
See Also ing enzymes and structural proteins, or a func-
tional RNA, including ribosomal RNA, transfer
▶ Codon RNA, small nuclear RNA, and other
▶ DNA non-translated RNAs. Gene expression involves
▶ Exon several sequential steps that are highly regulated,
▶ Genetic Code such as transcription, splicing, translation, and
▶ Genetics posttranslational modifications of proteins.
928 Gene Sequencing
See Also
Genetic Algorithms
▶ Gene
▶ Genetics Ann Nowé
▶ Protein Vrije Universiteit Brussel, Brussels, Belgium
▶ RNA
▶ Splicing
▶ Transcription Keywords
▶ Translation
Evolutionary algorithms
Synonyms
Gene Sequencing Darwinian approaches; Evolutionary algorithms
▶ DNA Sequencing
Definition
A genetic algorithm (GA) is a stochastic, parallel,

heuristic search algorithm inspired by the biolog-
Gene, Selfish ical model of evolution. It is used in computing to
find exact or approximate solutions to hard opti-
Stéphane Tirard mization and search problems.
des Techniques de Nantes, Nantes, France History
In his book on the origin of species, Charles

Gene selfish is a metaphoric expression coined Darwin brings forward variety, inheritance, and
for the first time in 1976 by the British biologist selection as important factors to explain the abil-
Richard Dawkins. Dawkins claimed that gene is ity of evolution to create the biological diversity
the “unit of selection” of the evolutionary profound in nature. This model was the source of
cess. In his conception, organisms are only sys- inspiration for genetic algorithms which embody
tems that permit to genes to reproduce. This idea a simpler version of the three previously men-
was central on the debate about the unity of tioned principles. John Holland is generally
selection that put in opposition Dawkins and accepted as the father of Genetic Algorithms.
S. J. Gould. Some other important contributors to the field in
Dawkins R., The Selfish Gene, Oxford Univer- the early years (1960s and 1970s) are
sity Press, 1976. I. Rechenberg, H.P. Schwefel, G. Box, and
L.J. Fogel.
Overview
General Circulation Model
Genetic algorithms belong to the family of Evo-
▶ GCM lutionary Algorithms (EA), which refers to a
class of stochastic, parallel search heuristics
Genetic Algorithms 929
inspired by the biological model of evolution. In order to simulate evolution, one needs the
The robustness of genetic algorithms has made equivalent of a chromosome as it is found in
it possible to apply them with success to many nature. A chromosome encodes genetic proper-
problems. This multitude of successes has also ties and can be modeled as a string of genes. The
led to the creation of many, often problem- interpretation of a chromosome is problem
specific, variations of the basic algorithm. The dependent. In the simplest scenario, the individ-
following section presents the standard genera- ual gene values (0 or 1) can be used to switch on
tional genetic algorithm. or off a specific feature. Alternatively, the bit
string can be used to encode a numeric value in
the form of a binary number.
Basic Methodology
Example
Population and Simulated Evolution Five chromosomes of eight binary genes each
A genetic algorithm uses a population of candi- (Table 1). G
date solutions. The population consists of a group
of individuals which are all potential solutions for
the problem one tries to solve. The use of a Evaluation: Fitness
collection of potential solutions is in sharp con- An evaluation function, also called the objective
trast with most classic search algorithms, such as or fitness function, has to be provided by the user
hill climbing, which typically only consider one to compute the score of an individual. The objec-
solution at a time. tive score plays a role similar to the environment
The population, together with the objective in an ecosystem as it provides a measure of the
(formulated by means of a fitness function), pro- quality of a solution for solving the given prob-
vides a context in which the individuals compete lem. In a classification problem, this can be the
for survival. Throughout the course of the simulated number of correctly classified instances, in which
evolution, the population will be subjected to differ- case the genetic algorithm will try to maximize
ent steps. These steps, which involve a fair amount the score.
of stochastic operations, will generate a new popu- The fact that genetic algorithms make very few
lation of candidate solutions. The whole process is assumptions concerning the objective function is
reiterated until a specified criterion is met. important. Classical methods like hill climbing use
The simulated evolution of the population information about the local gradient of the function
goes as follows: to guide the search process. This has the advantage
of pushing the search toward promising points in
1. Generation of the initial population the search space but also makes it susceptible to
2. Repeat until some user-specified criterion or a getting stuck in local optima. A genetic algorithm
certain number of generations have been does not use local gradient information concerning
exceeded: the objective function. Instead, it performs its
• Evaluation of the individuals by applying search process using several points in parallel in
the objective score (fitness function) the search space.
• ▶ Selection of one or more parent The fitness function, used to compute the fit-
individuals ness, converts the objective score into the fitness
• Application of genetic operators to build value in a range from 0 to 1. In the canonical
the next-generation population genetic algorithm, the fitness function, assuming
a maximization problem, is defined as the ratio
Genetic Algorithms, Table 1

Individual 01101010 11010111 00110100 01000101 11101111
930 Genetic Algorithms
between the objective score of an individual and virtually impossible for a genetic algorithm to
the total population objective score (TPOS) generate new candidate solutions, which implies
(in the example, this is 3.375). the search stagnates before reaching the optimal
solution.
Example Other often-used selection procedures are
A fitness function expressing an objective and tournament selection, where individuals are
the fitness of five candidates (Table 2). pairwise compared and the fittest individual is
selected with some given probability, and rank
selection, where individuals are ranked according
Selection to their fitness; the probability of selection is
Better individuals get more chance to survive due relative to the proportional rank.
to their intrinsically better qualities. Yet, as in
nature, less fit creatures do not necessarily disap- Example
pear, and fitter individuals may die by accident. Fitness proportional selection given the fitness
As mentioned in Darwin’s theory, diversity is of five candidates (Table 3).
needed for evolution to proceed. Thus, in order
to select candidate solutions (individuals) for the
next generation, a probabilistic selection proce- Genetic Operators
dure is required. Generally, part of the population To generate new solutions, a genetic algorithm
is replaced by selected individuals (on which any first selects one or more parent individuals using
combination of genetic operators is applied). The the provided selection procedure. It then applies
same individual may be selected multiple times. operators that either exploit the features present
One selection procedure, the fitness propor- in the selected individuals or explore new fea-
tional selection algorithm, selects individuals tures. Genetic algorithms mainly use three oper-
proportional to their relative fitness, meaning ators: (1) reproduction, (2) crossover, and
that the probability of an individual to be selected (3) mutation, which can be applied by themselves
equals the fitness as defined above. This can lead or in combination.
to premature convergence because the selection
pressure is too high. As a result, the diversity of Reproduction This is by far the simplest of all
the population is reduced too fast. It is then operators since it does not modify a parent

In our example, we are interested in finding solutions that have only strings of zeros or strings of ones as genes. Given
this objective,
Xa suitable objective function for individuals i, with g(i, k) denoting individual i’s kth gene, is
X
max g ði, k Þ; 8 gði, kÞ
f ðiÞ ¼ k¼1:::8
8
k¼1:::8
Individual 01101010 11010111 00110100 01000101 11101111

Objective score 0.5 0.75 0.625 0.625 0.875
TPOS = 3.375
Fitness 0.148 0.22 0.186 0.186 0.259

Individual 01101010 11010111 00110100 01000101 11101111
Fitness 0.148 0.22 0.186 0.186 0.259
Selection probability 0.148 0.22 0.186 0.186 0.259
Genetic Algorithms 931
individual. The individual is simply copied with- Many variants (such as the n-point operator)
out any change into the next generation. It is a exist and have proven their merits, mostly for
form of asexual reproduction. The idea of keep- specific problems and problem encodings.
ing a fraction of the population from one gener-
ation to the next is used in a more extreme and Example
less biologically faithful manner in what is called Two of the example individuals have been
elitism. Elitism systematically copies a selected for crossover. Using the one-point
predefined number of best individuals into the crossover operator, two new individuals are
next generation. In doing so, elitism completely generated for inclusion in the new population.
circumvents the normal, fitness proportional, Now, the next generation contains four indi-
selection scheme. viduals in total (Table 5).
Example
Two of the example individuals have been Mutation The purpose of the ▶ mutation oper- G
selected (individual 3 and 5) for reproduction. ator in genetic algorithms is to introduce diversity
In the example, for clarity, we apply only one in the gene pool. For the sake of simplicity,
operator to each selected (pair of) individual mutation is often implemented independently of
(s). This yields already two individuals for the crossover. Like the crossover operator, mutation
next generation (Table 4). is a nondeterministic process. Unlike crossover,
however, it does not have an exploitation charac-
ter but an exploring one. In fact, a mutation is
Crossover The crossover operator used by nothing more than a random perturbation.
genetic algorithms is the equivalent of sexual A mutation is performed in two steps, i.e., first,
reproduction in nature. It provides a randomized the position in the chromosome is chosen ran-
way of recombining the features of the two parent domly, and second, a new allele is chosen ran-
individuals in order to create two new ones and is domly. There can be a single or multiple
a simplified version of the recombination opera- mutations per individual.
tion that occurs between two DNA strands. The
crossover operator tries to exploit features that Example
are present in the parents and which have proven One individual is selected and mutated once.
useful. Crossover does not introduce new genetic Now, our next generation is completed and
variation. contains five individuals (Table 6).

Individual 01101010 11010111 00110100 01000101 11101111
Operator applied Reproduction Reproduction
New population 00110100 11101111

Individual 01101010 11010111 00110100 01000101 11101111
Operator applied One-point crossover
Swaps the genes, e.g., after gene 4: 0011|0100 & 0100|
0101 becomes 0011|0101 & 0100|0100
New population 00110100 11101111 00110101 01000100
932 Genetic Algorithms

Individual 01101010 11010111 00110100 01000101 11101111
Operator applied Mutation, gene 3
New population 00110100 11101111 00110101 01000100 11110111

Individual 00110100 11101111 00110101 01000100 11110111
Objective score 0.625 0.875 0.5 0.75 0.875
sum = 3.625
After selection and genetic operators have The schema theory predicts the discovery and
been applied, a new population is formed for survival of short combinations of features with
which the fitness can be calculated. This fitness above average fitness. Such a combination is
may, but not necessarily has to, improve over a called a building block. The building block
single generation. hypothesis is that the genetic search does not
proceed by merely exchanging and moving
Example genes around but by capitalizing on the building
In the example, the best objective score in the blocks that emerge in the population. The search
population has not improved, but the sum of process occurs through the juxtaposition of build-
objective score has, meaning that the popula- ing blocks. This reorganization is assumed to
tion improved compared to the previous gen- occur through operators like crossover.
eration (Table 7).
Repeating the process, the population will Applications

improve further, and better individuals will be
obtained via crossover and mutation. Genetic algorithms are usually applied to prob-
lems in the class NP, which are considered to be
very hard optimization problems (the time to find
Key Research Findings the solution scales in a nonpolynomial way with
the number of elements defining the problem).
Like any search algorithm, the genetic algorithm Some examples are scheduling, time-tabling,
comes with its set of parameters. The interaction traffic and shipment routing, engineering design,
between these parameters is often not well under- robotics, evolvable hardware, computer gaming,
stood. The encoding of the individuals and the and computer-aided molecular design. When a
fitness function determine the shape of the numerical optimization problem is considered,
so-called fitness landscape, which might have the strongly related class of algorithms called
many local optima. The more rugged the fitness evolution strategies can be a good candidate. An
landscape, the harder it is for the GA to find the evolutionary approach can also be adapted to
optimal solution. The most prominent theory to design computer programs, which is an approach
explain the problem-solving capabilities of generally referred to as Genetic Programming.
genetic algorithms is the schema theory
(Holland 1987). In his schema theory, Holland
explains the ability of a genetic algorithm to Future Directions
search a large space efficiently by modeling the
search process sampling hyperplanes in the The GA will perform better when interesting
search space rather than just points. building blocks are located near each other, so
Genetic Code 933
that the chance to be disrupted by crossover is not Synonyms

too large. When there are strong interactions
between genes that are located far from each Codon table
other, more advanced GAs should be used. This
is currently an important line of research within
the GA community. Definition
A genetic code defines how genetic information

See Also (stored within a polymerized ▶ nucleotide
sequence) translates into a ▶ protein
▶ Artificial Life (a polymerized ▶ amino acid sequence): specifi-
▶ Darwin’s Conception of the Origins of Life cally it is the set of rules that maps each possible
▶ Evolution, Biological triplet nucleotide subsequence (▶ codon) to its
▶ Mutation corresponding amino acid. For example, the G
▶ Natural Selection genetic code operating within the human nuclear
genome translates the mRNA codon containing
uridine-cytidine-adenosine into the amino acid ser-
ine. These rules emerge through base pairing
Back T, Fogel DB, Michalewicz Z (eds) (1997) Handbook between a codon and the anticodon of an appropri-
of evolutionary computation, vol 1. IOP Publishing, ate tRNA (“adaptor molecule”) that was previously
Bristol. ISBN 0-750-30392-1 charged with a specific amino acid by an appropri-
Gen M, Cheng R (1996) Genetic algorithms and engineer- ate aminoacyl tRNA synthetase enzyme (aaRS).
ing design. Wiley, New York
Goldberg DE (1989) Genetic algorithms in search, optimi-
zation and machine learning. Addison-Wesley, Boston
Haupt RL, Haupt SE (1998) Practical genetic algorithms. Overview
Wiley, New York
Holland JH (1987) Genetic algorithms and classifier systems:
foundations and future directions. In: Grefenstette JJ The phrase “genetic code” is widely associated
(ed) Proceedings of the second international conference with two misunderstandings that obscure its rel-
on genetic algorithms. Lawrence Erlbaum, Hillsdale evance to astrobiology.
Koza JR (1992) Genetic programming: on the program- First, popular media increasingly use the
ming of computers by means of natural selection. MIT
Press, Cambridge (here the individuals are programs or phrase “genetic code” to refer to the genetic
expressions). ISBN 0-262-11170-5 information contained within an organism. How-
Mitchell TM (1997) Machine learining. McGraw-Hill, ever, biologists use genetic code to refer to the
New York system of rules by which organisms translate
their genetic information into protein-based
metabolism. All known living systems require a
genetic code precisely because they partition bio-
Genetic Code chemistry into two different types of
polymers – one (nucleic acids) for storing genetic
Stephen Freeland information and another (proteins) for catalyzing
Astrobiology Institute, University of Hawaii the chemical reactions of metabolism. At present,
NASA, Honolulu, HI, USA it remains largely unknown whether we should
expect an independent origin of life to do like-
wise. Relevance to the origin, evolution, distribu-
Keywords tion, and future of life in the universe thus hinges
upon understanding how and why genetic coding
Amino acid; aaRS; Codon; mRNA; Protein; emerged as a central phenomenon of life on our
tRNA; Translation planet (Freeland 2009).
934 Genetic Code
a 2nd codon position b 2nd codon position

U C A G U C A G
UUU Phe UCU Ser UAU Tyr UAU Cys U UUU Phe UCU Ser UAU Tyr UAU Cys U
UUC Phe UCC Ser UAC Tyr UAC Cys C 3rd codon UUC Phe UCC Ser UAC Tyr UAC Cys C 3rd codon
U UUA Leu UCA Ser UAA Stop UAA Stop A position U UUA Leu UCA Ser UAA Stop UAA Trp A position
UUG Leu UCG Ser UAG Stop UAG Trp G UUG Leu UCG Ser UAG Stop UAG Trp G
1st codon position
CUU Leu CCU Pro CAU His CGU Arg CUU Thr CCU Pro CAU His CGU Arg
CUC Leu CCC Pro CAC His CGC Arg CUC Thr CCC Pro CAC His CGC Arg
C CUA Leu CCA Pro CAA Gln CGA Arg C CUA Thr CCA Pro CAA Gln CGA
CUG Leu CCG Pro CAG Gln CGG Arg CUG Thr CCG Pro CAG Gln CGG
AUU lle ACU Thr AAU Asn AGU Ser AUU lle ACU Thr AAU Asn AGU Ser
AUC lle ACC Thr AAC Asn AGC Ser AUC lle ACC Thr AAC Asn AGC Ser
A AUA lle ACA Thr AAA Lys AGA Arg A AUA Met ACA Thr AAA Lys AGA Arg
AUG Met ACG Thr AAG Lys AGG Arg AUG Met ACG Thr AAG Lys AGG Arg
GUU Val GCU Ala GAU Asp GGU Gly GUU Val GCU Ala GAU Asp GGU Gly
G GUC Val GCC Ala GAC Asp GGC Gly G GUC Val GCC Ala GAC Asp GGC Gly
GUA Val GCA Ala GAA Glu GGA Gly GUA Val GCA Ala GAA Glu GGA Gly
GUG Val GCG Ala GAG Glu GGG Gly GUG Val GCG Ala GAG Glu GGG Gly
Key
CGA Codon (nucleotide triplet)
Ser Amino acid translation product
Genetic Code, Fig. 1 Two examples of genetic codes: genetic code – in this case the one at work within the
(a) the standard genetic code at work in most genomes and mitochondria of Saccharomyces cerevisiae
present within LUCA. (b) An example of a nonstandard
Second, despite the continuing and wide- of correspondence between individual codons
spread use of the phrase “universal genetic and specific amino acids (e.g., Fig. 1). However,
code,” no such thing exists. Most known living this view is misleading because the process of
systems share in common a standard genetic code genetic coding is not exact: it is the outcome of
(Fig. 1a) that maps the 64 possible codons (i.e., a series of complex molecular interactions; thus a
each possible triplet combination of nucleotides tabulated summary is a statistical approximation.
bearing the bases uracil, thymine, ▶ adenine, or For a codon to be translated into an amino acid,
▶ cytosine) into 1 of 20 amino acid meanings. an appropriate tRNA must first be charged with
However, research over the past four decades has that amino acid by an appropriate aaRS enzyme.
revealed numerous nonstandard genetic codes at In some lineages this amino acid is then modified
work within naturally occurring genomes by further enzymes (e.g., see Wong 2005). The
(Knight et al. 2001) (e.g., Fig. 1b). Nonstandard charged tRNA must then compete against others
codes are widely distributed across diverse evo- in order to base-pair with the codon within the
lutionary sub-lineages, and all involve relatively molecular context of a ▶ ribosome (Betney
minor changes to the standard genetic code which et al. 2010). Errors are inherent to all of these
predominates within all three fundamental processes, and mismatching an amino acid to a
domains of life (Archaea, Bacteria, and given tRNA species (mischarging) or
Eukaryota). It therefore seems likely that the mismatching an incorrect codon to a correctly
standard genetic code was present within the charged tRNA species (misreading) causes devi-
▶ last universal common ancestor (LUCA) of ations from the code as written. While
extant life and that all nonstandard codes are mischarging of tRNAs appears comparatively
subsequent evolutionary diversifications. This rare (Ling et al. 2009), patterns and frequencies
shows that while genetic coding is fundamental of misreading vary widely between different lin-
to biology, it is not fully determined by physics or eages (Drummond and Wilke 2009). Thus, while
chemistry: it is an evolved and evolvable phe- the phrase “codon ambiguity” has been coined to
nomenon. Indeed, any genetic code under con- describe codons that are routinely translated into
sideration is usually represented as an exact table more than one amino acid at an unusually high
Genetic Code 935
frequency (Moura et al. 2009), a wealth of related probabilistically into a meaning drawn from a
research has shown that the fidelity of protein group of amino acids) (Fitch and Upper 1987).
translation depends to a significant (but widely A third major theme is that within the standard
varying) extent on “codon context,” where this is genetic code, the distribution of codons to amino
taken to mean the identity of neighboring codons acids appears very nonrandom in that amino acids
or of RNA motifs located outside of the codon with similar biochemical properties are assigned
itself (Buckingham 1994). This shows how the to codons that differ by only one of their three
precise rules of genetic coding can and do change nucleotides (Freeland et al. 2003). In general, this
during evolution, even for such a fundamental observation has been used to argue that natural
and complex feature as the standard genetic code. selection shaped the distribution of amino acid
Against this background, ongoing research is meanings to codons so as to reduce the harmful
aimed at understanding better the origin of effects of point mutation. More recently, it has
genetic coding (including the emergence of the been suggested that the same property may actu-
standard genetic code), the diversity of genetic ally enhance the efficiency of natural selection as G
codes found within present-day organisms (and it acts to adapt protein-coding genes within a
how these emerge), and the processes by which changing environment (Zhu and Freeland 2006).
biological engineers can manipulate a given These different major lines of research into the
genetic code to deviate from anything that occurs origin and emergence of a standard genetic code
naturally (e.g., by incorporating amino acids that do not necessarily compete as explanations. In
have never been part of any naturally occurring particular, far less attention has been given to
genetic code). the fundamental variables involved: for example,
Regarding the origin of genetic coding and the why are codons three nucleotides in length? Why
emergence of the standard genetic code, an does genetic coding use only four nucleobases
expansive and often speculative literature has and only 20 amino acids of the thousands of
converged around three major themes. First, that examples of each type of compound found in
genetic coding may have originated through biological systems (see Freeland (2009) for a
direct interactions between ▶ RNA motifs and review of ideas on these topics)? It remains
the amino acids that they encode (this may also entirely possible, though unknown, whether the
explain the codon assignments of the standard standard genetic code emerged as some sort of
genetic code, though to an unknown extent) optimal trade-off involving the steric affinities
(Yarus et al. Nov 2009). Second, it seems likely between mRNA motifs and amino acids, natural
that the standard genetic code emerged from a selection for an adaptive distribution of codon/
simpler genetic code comprising fewer codons amino acid assignments, and the growth of a
and/or fewer amino acids (more than 50 different standard amino acid alphabet from a near-infinite
versions of this broad idea are collated in set of possibilities.
Trifonov 2000). The most thoroughly developed Meanwhile, research over the past four
version, coined “genetic code coevolution,” decades has uncovered numerous slight variants
holds that we can see clues to the order by to the standard genetic code at work within the
which amino acids entered genetic coding by present-day biosphere (reviewed in Knight
comparing the codon assignments of the standard et al. 2001). Variations typically involve the
genetic code with conserved metabolic pathways reassignment of one or more codons to a different
of modern organisms, in which chemically com- amino acid meaning from that found in the stan-
plex amino acids are still formed through biosyn- dard genetic code (e.g., within the human mito-
thetic manipulation of their simpler amino acid chondrial genome, the codon AUA corresponds
precursors (Wong 2007). An alternative view is not to isoleucine but to methionine). Many occur
that all codons and amino acids were present within the highly reduced genomes of mitochon-
from the start, but coding was originally highly dria (e.g., eight codons have changed amino acid
ambiguous (i.e., groups of codons were translated assignments within the genetic code that operates
936 Genetic Code
on the mitochondrial genome of yeast, Saccharo- recent origin of this research, its rapid success,
myces cerevisiae – Fig. 1b), but others exist and the inherent economic and biomedical poten-
within free-living microbes. Nonstandard codes tial, significant future progress seems likely. In
also include examples that use one of two non- particular, scientists elsewhere have successfully
standard amino acids (▶ selenocysteine and incorporated additional bases into DNA (e.g., see
pyrrolysine) (Yuan et al. 2010). Although some Geyer et al. 2003). This offers the potential for a
maintain that these amino acids are not truly combination of the two technologies to greatly
encoded in the same sense as the standard enlarge the complexity of the genetic code, pro-
20, owing to unusual features associated with ducing proteins that have no analogs within
the process of their translation, this argument nature, without disrupting current genomes and
fails under careful examination (Freeland 2009). their translation into metabolism.
Owing to recent discoveries of enormous, largely
uncharacterized microbial biodiversity (Fierer
et al. 2007), it seems likely that current science
is far from knowing the full range of nonstandard See Also
codes that currently exist within nature. This may
well include the presence of additional coded ▶ Activated Nucleotide
amino acids (although bioinformatics analyses ▶ Adenine
of the world’s repositories of protein-coding ▶ Amino Acid
sequence data suggest that further such ▶ Anticodon
discoveries are unlikely to occur within species ▶ Aptamer
whose genes have been sequenced so far) ▶ Biopolymer
(Freeland 2009). Debate still surrounds the mech- ▶ Cytosine
anisms by which codons become reassigned to ▶ Gene
new amino acid meanings (e.g., see Sengupta ▶ Genetics
et al. 2007), but a large quantity of empirical ▶ Guanine
data shows that mistakes routinely occur in the ▶ L-Amino Acids
process by which tRNAs bond with their ▶ Last Universal Common Ancestor
cognate codons. In some cases, this mismatching ▶ Nucleic Acid Base
reaches levels at which a codon is declared ▶ Nucleotide
“ambiguous” with respect to its amino acid ▶ Peptide
meaning (e.g., Moura et al. 2009). More gener- ▶ Protein
ally, evidence is growing that many features of an ▶ Ribosome
mRNA sequence can influence the accuracy with ▶ RNA
which a codon is translated. As an evolutionary ▶ Selenocysteine
route to codon reassignment, this is consistent ▶ Thymine (T)
with the finding that the two nonstandard amino ▶ Translation
acids to have been incorporated into coding ▶ Uracil (Ura)
are typically associated with mRNA motifs
that lie outside of the codon itself (reviewed in
Yuan et al. 2010).
More recently, researchers have begun to Betney R, de Silva E, Krishnan J, Stansfield I (2010)
demonstrate success in manipulating the rules of Autoregulatory systems controlling translation factor
genetic coding within the laboratory. In particu- expression: thermostat-like control of translational
lar, the emerging science of synthetic biology has accuracy. RNA 16:655–663, [Epub ahead of print],
PMID: 20185543
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into the standard genetic code within model thesis: enhancements of the genetic code. Biochimie
organisms (Xie and Schultz 2006). Given the 76(5):351–354, Review
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Nat Rev Genet 2:49–58 a linear map on which the distances are expressed
Ling J, Reynolds N, Ibba M (2009) Aminoacyl-tRNA as frequency of crossing-over or percent
synthesis and translational quality control. Annu Rev ▶ recombination. Genetic distance in the map
Microbiol 63:61–78, Review, PMID: 19379069 has no physical units: Its unit – m.u or
Moura GR, Carreto LC, Santos MA (2009) Genetic code
ambiguity: an unexpected source of proteome innova- centiMorgan – corresponds to 1 % of recombina-
tion and phenotypic diversity. Curr Opin Microbiol tion. Genetic distance describes the tendency of
12(6):631–637, Epub 21 Oct 2009 Review two given genes on the same chromosome to be
Sengupta S, Yang X, Higgs PG (2007) The mechanisms of inherited together: If they are physically close to
codon reassignments in mitochondrial genetic codes.
J Mol Evol 64(6):662–688, Epub 29 May 2007 each other, they will tend to stay together with-
Trifonov EN (2000) Consensus temporal order of amino standing the recombination events produced dur-
acids and evolution of the triplet code. Gene ing meiosis, a form of cell division in eukaryotes
261(1):139–151 essential for sexual reproduction. In turn, genes
Wong JT (2005) On the formation of Asp-tRNA(Asn) by
aspartyl-tRNA synthetases. Bioessays 27:1309 far away from one another will have a long
Wong JT (2007) Question 6: coevolution theory of the genetic distance since they show a higher proba-
genetic code: a proven theory. Orig Life Evol Biosph bility to be separated as a result of recombination.
37(4–5):403–408 Thus, the physical proximity on a same chromo-
Xie J, Schultz PG (2006) A chemical toolkit for proteins-
an expanded genetic code. Nat Rev Mol Cell Biol some explains the correlation with the degree of
7:775–782 genetic linkage evidenced in the crossing exper-
Yarus M, Widmann JJ, Knight R (2009) RNA-amino acid iments. Overall, a ▶ genome has as many linkage
binding: a stereochemical era for the genetic code. groups as chromosomes. For example, humans
J Mol Evol 69(5):406–429
Yuan J, O’Donoghue P, Ambrogelly A, Gundllapalli S, have 24 linkage groups: the
Lynn Sherrer R, Palioura S, Simonović M, Söll 22 autosomes – chromosomes that contain
D (2010) Distinct genetic code expansion strategies genetic information available to both
938 Genetics
sexes – plus the sex chromosomes X and Y. Gene Applied branches of genetics rely on the
mapping has important applications, including established genetic principles and are used in
the production of the first drafts during a genome medicine, agriculture, and biotechnology.
sequencing project.
Overview
See Also
Early naturalists essentially observed the herita-
ble traits of animals or plants in an empirical
▶ Chromosome
manner, based on a long tradition of experimental
▶ DNA Sequencing
breeding dating back several centuries. This
▶ Gene
approach was followed by pedigree analysis, by
▶ Genome
which the inheritance of some specific traits was
▶ Genomics
traced in all members of a family line. For exam-
▶ Recombination
ple, human pedigrees traced the inheritance of
certain diseases such as polydactylism, hemo-
philia, or albinism. G. J. Mendel, considered the
father of genetics, described in 1866 the basis of
Genetics inheritance of traits from one generation to the
next. At the beginning of the twentieth century,
Carlos Briones after the rediscovery of Mendel’s work and fol-
Centro de Astrobiologı́a (CSIC/INTA), Consejo lowing an increasing interest in cytology and
Superior de Investigaciones Cientı́ficas, Madrid, biochemistry, the term “genetics” was coined in
Spain 1905 by W. Bateson to describe the study of
heredity and variation (Bateson 1907). The field
of population genetics came into being in the
Keywords 1920s to study the distribution of ▶ genes in
biological populations and their change under
DNA; Flow of genetic information; Gene; the influence of evolutionary processes. It was
Genome; Inheritance; Molecular biology; based on the synthesis of the principles of Men-
Protein; RNA delian genetics with Darwinian natural selection
(Sturtevant 2001).
The observation that DNA is the genetic mate-
Definition rial in all ▶ cells and the discovery of the struc-
ture of DNA as a double helix inaugurated the era
Genetics is the discipline of Biology which stud- of molecular genetics in the decade of the 1950s.
ies inheritance, variation, function, and the phys- The investigation within this field has shown that
ical nature of the genetic material. Historically, all cells and a fraction of ▶ virus families have
genetics has proceeded along several major path- DNA genomes, while RNA is the genetic mate-
ways including experimental breeding, pedigree rial of other viruses and all ▶ viroids. The overall
analysis, population genetics, cytogenetics, flow of genetic information in cells and viruses
molecular genetics, and genetic engineering. can be represented as DNA$RNA ! Proteins,
Among them, molecular genetics is mainly as schematized in Fig. 1. The spectacular pro-
concerned with the molecular bases of the inher- gress produced in molecular biology opened the
itance and variation of the ▶ genotypes and possibility to introduce tailored modifications in
investigates the relationship between the the genetic material of organisms, which gave
information-bearing macromolecules – DNA rise to genetic engineering in the 1970s (Alberts
and RNA – and the proteins encoded by them. et al. 2002; Lewin 2008).
Genetics 939
Genetics,
Fig. 1 Schematic
representation of the flow DNA
of genetic information.
Cellular processes are
shown by solid arrows, Replication
while those found only in
viruses are depicted as Transcription Retro-
dotted arrows
transcription
RNA
Replication
Translation G
Protein
Since 1990, the advances in DNA sequencing See Also

and bioinformatics opened the possibility to
study whole genomes of organisms as well as ▶ Cell
their set of encoded products, thus initiating the ▶ Evolution, Biological
current era of genomics and proteomics. The ▶ Gene
publication of the first draft of the human ▶ Genetics, History of
▶ genome in 2001 and its relevance for modern ▶ Genome
medicine has expanded broadly the subfield of ▶ Genotype
human molecular genetics (Strachan and Read ▶ Metagenome
2004; Watson et al. 2008). In turn, recent ▶ Proteome, Proteomics
advances in biotechnology allowed the study of ▶ Replication (Genetics)
the “▶ metagenome” of communities containing ▶ Transcription
different species (e.g., Venter et al. 2004), as well ▶ Translation
as the analysis of genetic material from extinct ▶ Viroid
organisms, in the interdisciplinary field of ▶ Virus
paleogenetics (e.g., Poinar et al. 2006).
In summary, the tools and developments of
genetics are currently integrated into all biologi-
cal disciplines, including origin and evolution of Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter
life, taxonomy, microbiology, virology, molecu- P (2002) Molecular biology of the cell. Garland, New
lar biology, biochemistry, biophysics, biotech- York
nology, cytology, physiology, and development. Bateson W (1907) The progress of genetic research. In:
Wilks W (ed) Report of the third 1906 international
Additionally, advances in genetics have perme- conference on genetics: hybridization (the cross-
ated other fields such as medicine, forensics, psy- breeding of genera or species), the cross-breeding of
chology, sociology, paleontology, and varieties, and general plant breeding. Royal Horticul-
astrobiology. tural Society, London
940 Genetics, History of
Lewin B (2008) Genes IX. Jones and Bartlett, Boston These empirical observations did not constitute
Poinar HN, Schwarz C, Qi J et al (2006) Metagenomics to a science of heredity, the systematic study of the
paleogenomics: large-scale sequencing of mammoth
DNA. Science 311:392–394 transmission of characters through generations.
Strachan T, Read AP (2004) Human molecular genetics. The father of the science of heredity is Gregor
Garland, New York J. Mendel, who in the garden of the monastery of
Sturtevant AH (2001) A history of genetics. Cold Spring Brno discovered the so-called laws of heredity.
Harbor Laboratory Press, New York
Venter JC, Remington K, Heidelberg JF, Halpern AL, The work of Mendel was ignored by his contem-
Rusch D, Eisen JA, Wu D, Paulsen I, Nelson KE, poraries (including Charles R. Darwin), but was
Nelson W, Fouts DE, Levy S, Knap AH, Lomas rediscovered 35 years later, on the eve of the
MW, Nealson K, White O, Peterson J, Hoffman J, twentieth century, by Hugo de Vries, Carl
Parsons R, Baden-Tillson H, Pfannkoch C, Rogers Y,
Smith HO (2004) Environmental genome shotgun Correns, and Erich von Tschermak.
sequencing of the Sargasso sea. Science 304:66–74 Mendel is the emblematic example of a “pre-
Watson J, Baker T, Bell S, Gann A, Levine M, Losick cursor,” a man or woman who anticipates the
R (2008) Molecular biology of the gene. Benjamin future progress of knowledge. Historians have
Cummings, Menlo Park
done much to dispel the legend surrounding Men-
del and his discovery. He was not a monk isolated
in his monastery, but a recognized scientist, in
contact with other scientists of his time, and well
Genetics, History of aware of the most recent scientific developments.
His project was not the establishment of the laws
Michel Morange of heredity, but the search for an answer to a
Centre Cavaillès, USR 3308 CIRPHLES, Ecole practical question faced by all gardeners and
normale supérieure, Paris Cedex 05, France breeders: why are some hybrids stable and others
unstable? And when the hybrids are not stable,
how do they return to the parental types? He had
Keywords the merit to look for (and discover) quantitative
laws of transmission of characters. But he had no
Chromosome; Development; DNA; Gene; clear idea of “what” transmitted these characters
Genetic code; Genome; Mendel; Phenotype- from one generation to another.
genotype The 35 years which separated Mendel from
the rediscovery of his laws was necessary to
convince biologists of the existence and impor-
History tance of the mechanisms of heredity. In 1859,
simultaneously but independently of Mendel’s
It is quite impossible to pinpoint the origin of work, Darwin proposed his theory of evolution,
▶ genetics. The resemblances, but also the differ- which called for the existence of a mechanism of
ences, between children and their parents transmission through generations of the sponta-
(or grandparents), as well as the experience of neous variations screened by ▶ natural selection.
breeders with animals, represent common ances- Ten years later, Darwin proposed his model of
tral knowledge. The evidence that characteristics, pangenesis. This model was neither original nor
but also potentialities, were transmitted from one based on experimental evidence. But it triggered
generation to the next was widely accepted. the work of many biologists, including August
These spontaneous “prescientific” observations Weismann and Hugo de Vries, who proposed
coexisted with the theory of impregnation, the alternative models of heredity. In parallel and in
conviction that characteristics acquired during relation with cytological observations on the
life could be transmitted, and the idea that the nucleus, the chromosomes, and the mechanisms
thoughts and imagination of the mother might of fertilization, these authors elaborated what the
imprint their shapes on the future newborn. historian of biology Garland E. Allen has
Genetics, History of 941
described as corpuscular and mechanistic models and embryology and the autonomy of the former.
of heredity (Allen 2000). Minds are now ripe for Indeed, the nature of genes and the mechanisms
the rediscovery of Mendel’s laws. by which genes control the formation of charac-
The first 10 years of the science of genetics ters were difficult scientific questions, experi-
(1900–1910) was necessary to separate charac- mental approaches to which were still lacking.
ters from genes or, as Wilhelm Johannsen said, This abstract nature of the gene did not prevent
▶ phenotype from ▶ genotype and to propose the wedding between genetics and Darwinism
that the chromosomes were the bearers of the which occurred in the 1930s through the work
genes (a hypothesis made by W. S. Sutton and of John Haldane, Sewall Wright, and Ronald
Th. Boveri in 1902). A brilliant confirmation of A. Fisher. They elaborated the sophisticated
this hypothesis and the choice of an appropriate mathematical models of population genetics,
experimental system (the fruit fly Drosophila which provided a precise definition of ▶ fitness,
melanogaster) were due to Thomas H. Morgan and demonstrated that mutations of small ampli-
and his small group of collaborators working at tude and low selective pressures were sufficient G
Columbia University in New York (Allen 1978). to drive the transformation of organisms. The
Sturtevant and Morgan rapidly demonstrated that modern synthesis, the new theory of evolution,
the genes present on the same chromosomes were emerged in the 1940s from the transfer of genetic
not always transmitted together. They interpreted studies done in the laboratory to fieldwork
this observation thanks to the 1909 discovery by (Theodosius Dobzhansky), from an elaboration
the Belgian cytologist R. A. Janssens of the of complex mechanisms of speciation (Ernst
recombination of the chromosomes during meio- Mayr), and from an interpretation of paleontolog-
sis (the cell division mechanism by which germ ical data in the light of the models of population
cells eliminate half of their chromosomes). This genetics (George Simpson).
explanation allowed the drawing of genetic maps: Meanwhile, the abstract notion of the gene
the probability of genes being separated by chro- was progressively replaced by a pure chemical
mosome breaks is proportional to the physical description. In the 1930s, cytochemical studies
distance separating these genes on the demonstrated that ▶ DNA and proteins were the
chromosome. main constituents of chromosomes. In 1944,
Rapid progress was made in genetics, not only Oswald Avery and his collaborators produced
with Drosophila but also with maize and humans. strong arguments in favor of a major role of
Despite this intense effort of genetic mapping, the DNA in the process of heredity. The double-
questions of the nature of the ▶ gene and of the helix model proposed by Jim Watson and Francis
mechanisms by which genes were responsible for Crick paved the way to the explanation of how
the formation of characters not only were not the DNA molecule was faithfully replicated and
addressed but were ignored. In his Nobel lecture suggested that the order of bases in DNA was
(1934), Morgan still stated that defining the probably the basis of genetic information. At the
nature (material or not) of the genes would not beginning of the 1960s, the genetic code was
alter the research program of geneticists. deciphered, demonstrating how the nucleotide
Geneticists’ apparent lack of interest in the ▶ sequence of DNA controlled the order of
nature and mechanism of action of genes proba- amino acids in proteins and therefore their struc-
bly has its roots in the empiricist philosophical tures. Genes had been reduced to DNA. The iden-
tradition which permeated the minds of many tification of the gene with an object seemed
scientists and biologists at the beginning of the complete.
twentieth century. Establishing the laws by which But the situation rapidly deteriorated. The dis-
characters were transmitted through generations, covery of regulatory sequences, located upstream
and ignoring the mechanisms by which they were of the coding regions of the genes, initiated a
reproduced at each generation, was also a strat- process of structural deconstruction of the gene.
egy to maintain the separation between genetics These sequences cannot belong to the gene, since
942 Genome
they can be shared by different genes. Neverthe- ▶ Genotype

less, they are necessary for the proper activity ▶ Monod’s Conception on the Origins of Life
(expression) of the gene. Other difficulties ▶ Natural Selection
emerged: the existence of split genes in eukary- ▶ Phenotype
otes, the capacity for a DNA sequence to generate ▶ Protein
different messenger RNAs and proteins, the ▶ RNA
editing process, and the existence of DNA ▶ Sequence
sequences coding in different frames for totally
distinct proteins. These were some of the obser-
vations preventing a simple definition of
the gene. Allen GE (1978) Thomas Hunt Morgan: the man and his
The same difficulties also emerged at the func- science. Princeton University Press, Princeton
tional level. Initially, it was admitted that a gene Allen GE (2000) The reception of Mendelism in the
coded for an enzyme, a ▶ protein with a well- United States, 1900–1930. C R Acad Sci III Sci Vie
323:1081–1088
determined chemical function. Other protein Keller EF (2000) The century of the gene. Harvard Uni-
functions were progressively discovered: ligand versity Press, Cambridge
receptors, transcription factors, adaptors, Morange M (1998) A history of molecular biology. Har-
etc. These new functions only find their meaning vard University Press, Cambridge
Sturtevant AH (1965) A history of genetics. Harper and
at a more integrated level, by conjunction with Row, New York
other proteins. Many genes participate in the for-
mation of one single character. This was not a
real surprise for geneticists, but it found its expla-
nation in the progressive description of these
molecular actors. In most cases, the expression Genome
“gene for” has no meaning!
The present situation is not simple, and efforts Carlos Briones
to provide a general definition of a gene have Centro de Astrobiologı́a (CSIC/INTA), Consejo
failed. Is the solution to be found in the replace- Superior de Investigaciones Cientı́ficas, Madrid,
ment of the single notion of gene by different Spain
well-defined concepts (Keller 2000)? Many
have argued for this and have even proposed
these new concepts. But the notion of gene is Keywords
still alive and well. Its vagueness allows this
“boundary object” to circulate between different Chromosome; DNA; Gene; Genetic information;
disciplines and to fertilize each of them. RNA; Sequencing project
See Also Definition
▶ Darwin’s Conception of the Origins of Life The genome is the total amount of the genetic
▶ DNA information of a ▶ species. It can also be defined
▶ Evolution, Biological as the basic haploid chromosome set of a species.
▶ Fitness The term “genome” has also been used to refer to
▶ Gene the full hereditary information of an organism.
▶ Genetics Nevertheless, the term ▶ “genotype” is preferred
▶ Genetic Map for individual organisms, since the “genome” of a
▶ Genome given species is in practice an average or consen-
▶ Genomics sus of the genetic information of a number of
Genome 943
Genome, Table 1 Sequenced genomes from representative viroids, viruses, and cellular organisms from the three
domains of life
Species Genomic material Genome size (nt) Release year
Viroid
Avocado sunblotch viroid (ASBVd) ssRNA 2.5 102 1981
Potato spindle tuber viroid (PSTVd)a ssRNA 3.6 102 1978
Virus
Hepatitis delta virus (HDV)b ssRNA 1.7 103 1987
Duck circovirus ssDNA 2.0 103 2005
Hepatitis B virus (HBV) ss/dsDNA 3.2 103 1987
Bacteriophage MS2a ssRNA 3.6 103 1976
Tobacco mosaic virus (TMV) ssRNA 6.4 103 1983
Simian rotavirus SA11c dsRNA 1.9 104 1990
Bacteriophage KVP40 dsDNA 2.4 105 2003
Mimivirus dsDNA 1.2 106 2004
G
Bacteria
Buchnera aphidicola strain Cc dsDNA 4.2 105 2006
Haemophilus influenzaea dsDNA 1.8 106 1995
Bacillus subtilis dsDNA 4.2 106 1997
Escherichia coli strain K-12 dsDNA 4.6 106 1997
Chitinophaga pinensis dsDNA 1.8 107 2009
Archaea
Nanoarchaeum equitans dsDNA 4.9 105 2003
Methanocaldococcus jannaschiia dsDNA 1.7 106 1996
Pyrococcus furiosus dsDNA 1.9 106 1999
Sulfolobus solfataricus dsDNA 2.7 106 2009
Haloarcula marismortui dsDNA 4.3 106 2004
Eukarya
Saccharomyces cerevisiaea dsDNA 1.2 107 1996
Plasmodium falciparum dsDNA 2.3 107 2002
Vitis vinifera dsDNA 4.9 108 2007
Homo sapiens dsDNA 3.2 109 2001
Monodelphis domestica dsDNA 3.5 109 2007
a
First complete genome sequenced within each group
b
Defective virus
c
Segmented viral genome
individuals belonging to that species. Thus, the circular topologies, and they are composed of
genome of one species does not capture the either a single DNA or RNA molecule or a set
genetic polymorphisms within it. of different ones, each of them carrying a frag-
ment of the whole genetic information. Genome
sizes vary from less than 300 nucleotides (nt) in
Overview the smallest viroids to up to 1011 base pairs (bp) in
certain vertebrate animals and higher plants.
All cellular genomes are composed of double- Some species have haploid genomes with a single
stranded (ds) ▶ DNA, while ▶ viruses can have copy of their genetic information, while others
DNA or ▶ RNA genomes as both single-stranded harbor multiple copies of the information in their
(ss) and ds molecules. In turn, all ▶ viroids show diploid, triploid, or polyploid genomes (Lewin
ssRNA genomes. Genomes can have linear or 2008; Gibson and Muse 2009).
944 Genome
Cellular genomes show one or more biomedicine and biotechnology, the advances in
▶ chromosomes – molecules of dsDNA with a genomics have largely influenced our current
certain degree of structural organization – view of different disciplines of biology (Snustad
together with, in most microorganisms, extra- and Simmons 2010).
chromosomal elements such as ▶ plasmids,
viruses, or transposons. In eukaryotes, apart
from the chromosomes composing their nuclear
genome, two organelles – mitochondria and See Also
chloroplasts – harbor dsDNA genomes contribut-
ing to the overall genome of the species. ▶ Bioinformatics
Genomes contain both the coding ▶ Cell
▶ genes – nucleic acid sequences that will be ▶ Chromosome
translated into proteins – and the noncoding ▶ DNA
genetic information. The latter includes the ▶ DNA Sequencing
rRNA, tRNA, and other small RNA genes, ▶ Gene
as well as different kinds of regulatory sequences, ▶ Genetic Map
structural elements, and sequences whose ▶ Genetics
function is at present unknown. Comparative ▶ Genotype
genomic methods have been widely used to iden- ▶ Metagenome
tify genes and regulatory regions in different ▶ Plasmid
species, as well as to trace patterns of ▶ RNA
genome evolution (Gibson and Muse 2009; ▶ Species
Nielsen et al. 2010). ▶ Viroid
Genome projects are aimed at determining the ▶ Virus
sequence of the entire genome of one species.
Upon completion of a sequencing project, the References and Further Reading
physical or ▶ genetic map of the genome and
the inventory of annotated genes and regulatory Collins F (2010) Has the revolution arrived? Nature
regions become available. The first complete 464:674–675
Fiers W, Contreras R, Duerinck F, Haegeman G,
genome sequenced belonged to an RNA virus, Iserentant D, Merregaert J, Min Jou W, Molemans F,
the bacteriophage MS2 (Fiers et al. 1976), Raeymaekers A, Van den Berghe A, Volckaert G,
followed by the DNA genome of the bacterio- Ysebaert M (1976) Complete nucleotide sequence of
phage PhiX174 (Sanger et al. 1977). Since then, bacteriophage MS2 RNA: primary and secondary
structure of the replicase gene. Nature 260:500–507
thousands of complete viral and cellular genomes Gibson G, Muse SV (2009) A primer of genome science.
have been already sequenced or are in progress. Sinauer Associates, Sunderland
Table 1 shows representative examples of Lander E, Linton L, Birren B, International Human Gene
sequenced genomes. The sequencing projects of Sequencing Consortium et al (2001) Initial sequencing
and analysis of the human genome. Nature
long and complex genomes, such as those from 409:860–921
higher plants and animals, require years of joint Lewin B (2008) Genes IX. Jones and Bartlett, Boston
experimental and bioinformatic work. Among Nielsen CB, Cantor M, Dubchak I, Gorton D, Wang
them, the Human Genome Project produced the T (2010) Visualizing genomes: techniques and chal-
lenges. Nat Methods 7:S1–S11
first draft of the complete genome of our species Sanger F, Air GM, Barrell BG, Brown NL, Coulson AR,
in 2001 (Lander et al. 2001; Venter et al. 2001). Fiddes CA, Hutchison CA, Slocombe PM, Smith
This achievement was followed by a rapid accel- M (1977) Nucleotide sequence of bacteriophage phi
eration in genome science and ▶ DNA sequenc- X174 DNA. Nature 265:687–695
Snustad DP, Simmons MJ (2010) Principles of genetics,
ing techniques, what has opened the door to a 5th edn. Wiley, New Jersey
future genome-based, personalized medicine Venter JC, Adams M, Myers E et al (2001) The sequence
(Collins 2010). Apart from their applications in of the human genome. Science 291:1304–1351
Genome, Minimal 945
It is not possible to specify a universal

Genome, Minimal minimal genome. First, it needs to be tied to a
particular level of biological organization.
Rosario Gil The increasing knowledge on bacterial
Institut Cavanilles de Biodiversitat i Biologia genomes and the simplicity of prokaryotic cells
Evolutiva, Universitat de València, Paterna makes bacterial cells a suitable choice. Second,
(València), Spain metabolism is extremely variable and highly
dependent on the environment. Therefore, a
plethora of minimal genomes, composed by a
Keywords universal informational core plus a variable met-
abolic gene set, can be envisaged (Gil
Core genome; Minimal cell; Minimal gene set et al. 2004).
Experimental and computational methods
have been used to define the core of a minimal G
Synonyms bacterial genome (Gil 2013). Experimentally,
essential genes are determined based on indirect
Minimal gene set evidence from systematic and genome-wide inac-
tivation or inhibition of each individual gene
present in a genome (compiled in http://www.
Definition essentialgene.org/). Computational comparative
genomics has been also extensively employed,
A minimal genome was defined in 1999 by assuming that genes shared between distant
Arcady Mushegian as a genome containing the organisms are likely to be essential. Naturally
smallest number of genetic elements sufficient to reduced genomes from bacteria with a host-
build a modern-type free-living cellular organism associated lifestyle have been used for compari-
(Mushegian 1999). Consequently, the concept of sons because they must be closer to a minimal
minimal genome is fundamentally related with genome (Gil et al. 2003; Mushegian and Koonin
the definition of a ▶ minimal cell. 1996). These studies showed the importance of
considering that essential functions can be
performed by alternative and unrelated
Overview (non-orthologous) proteins. Comparative ana-
lyses only retrieve genes involved in functions
Cells usually cannot import functional proteins or for which there is no alternative in nature (e.g.,
RNAs, so they rely on their own gene products to the complex translational machinery), while a
perform all essential functions. Once the essential minimal genome must also include all genes
functions have been identified, it is possible to essential to maintain metabolic homeostasis (Gil
delineate the gene set needed to perform them. et al. 2004).
Most attempts to define a minimal genome focus A third approach to a minimal genome
on protein-coding genes, ignoring functional searches for the biochemical and modular
RNAs, regulatory and other noncoding sequences, description of well-defined pathways needed to
and the organization of these elements on chromo- perform all essential functions (Jewett and For-
somes. Therefore, they only represent the core of a ster 2010). Although some important challenges
hypothetical minimal modern-cell genome. Sev- still need to be faced, this approach allows a
eral estimates indicate that such minimal genome function-by-function debugging to reach self-
core must contain at least about 200 genes, replication, and it seems a good start to approach
although some authors point out that additional the goal of synthesizing a minimal genome able
genes might be necessary to improve cell effi- to sustain an artificial minimal cell, the ultimate
ciency (Jewett and Forster 2010). goal of synthetic biology.
946 Genomics
See Also the completion of the first bacterial genome in

1995, the first eukaryotic genome in 1996, and the
▶ Cell, Minimal release of the first draft of the human genome in
▶ Endosymbiosis 2001. Different current subfields of genomics
include structural genomics, functional geno-
mics, metagenomics, pharmacogenomics, and
comparative genomics.
Gil R (2014) The minimal gene-set machinery. In: Meyers
RA (ed) Encyclopedia of molecular cell biology and
molecular medicine. Wiley-Blackwell, Weinheim, pp.
1–36 See Also
Gil R, Silva FJ, Zientz E, Delmotte F, Gonzalez-Candelas-
F, Latorre A, Rausell C, Kamerbeek J, Gadau J, ▶ Bioinformatics
Holldobler B, van Ham RCHJ, Gross R, Moya
▶ DNA Sequencing
A (2003) The genome sequence of Blochmannia
floridanus: comparative analysis of reduced genomes. ▶ Genetics
Proc Natl Acad Sci U S A 100:9388–9393 ▶ Genome
Gil R, Silva FJ, Pereto J, Moya A (2004) Determination of ▶ Metagenome
the core of a minimal bacterial gene set. Microbiol Mol
▶ Proteome, Proteomics
Biol Rev 68:518–537
Jewett MC, Forster AC (2010) Update on designing and
building minimal cells. Curr Opin Biotechnol
21:697–703
Mushegian A (1999) The minimal genome concept. Curr
Opin Genet Dev 9:709–714 Genotype
Mushegian AR, Koonin EV (1996) A minimal gene set
for cellular life derived by comparison of complete
bacterial genomes. Proc Natl Acad Sci U S A 93: Susanna Manrubia
10268–10273 Systems Biology Program, Centro Nacional de
Definition
Genomics
The genotype is the genetic makeup of an organism,
Carlos Briones its full hereditary information. The genotype of an
Centro de Astrobiologı́a (CSIC/INTA), Consejo organism is a characteristic fingerprint. Genotype is
Superior de Investigaciones Cientı́ficas, Madrid, different from ▶ genome (usually describing a spe-
Spain cies), which is a representative average over indi-
viduals. For example, a viral ▶ quasispecies
contains a particularly large number of different
Definition genotypes, though the ▶ virus is characterized by
a precise genome. In diploid organisms, the geno-
Genomics is the field of ▶ genetics which studies type is the pair of alleles inherited for each partic-
the structure, organization, and function of entire ular gene. The expression of the genotype in a given
genomes. This global and integrative approach environment is the ▶ phenotype of an organism.
was initiated in the decade of the 1980s, thanks
to the advances in automatic ▶ DNA sequencing
and the availability of new bioinformatic tools for See Also
analyzing increasingly large amounts of genetic
information. Four milestones in genomics were ▶ Genetics
the sequencing of the first viral genome in 1976, ▶ Genome
Geochronology 947
▶ Mutation
▶ Phenotype Geochronology
▶ Quasispecies
▶ Virus Francis Albarède
France
Genus Keywords
Ricardo Amils Age; Absolute age; Ion microprobe; Isochron;

Departamento de Biologia Molecular, Isotope; Radioactivity; Radioactive decay;
Universidad Autónoma de Madrid, Madrid, Mass spectrometer; Extinct radioactivity; Solar
Spain System G
Definition
Synonyms
Genus is a taxonomic group of related
Age measurement; Radiochronology
▶ species sharing one or more major properties.
The term comes from the Latin genus meaning
“descent, family, type, and gender,” cognate with
Definition
Greek: genoB-genos, “race, stock, and kin.”
A genus is defined through the characterization
Geochronology is the scientific field dedicated to
of different species. In order for a genus to be
providing absolute ages of geological and plane-
descriptively useful, it must have monophyly,
tary material. It is based on the measure of the
reasonable compactness, and distinctness. In the
accumulated isotopes produced by the ▶ radioac-
binomial nomenclature used in biology, the
tivity of some long-lived ▶ nuclides. The radio-
genus is used as the first word of a scientific
activity of now extinct nuclides can be used to
name. The genus name is always capitalized and
date the early evolution of the solar system. Most
italicized. In the hierarchy of the binomial classi-
of the techniques involved in this field are based
fication system, genus comes above species and
on ▶ mass spectrometry.
below family. In the past, the taxonomic
adscription of organisms was based on pheno-
typic properties, but since the introduction of
molecular biology, techniques allowing the use History
of genotypic properties have facilitated the estab-
lishment of phylogenetic relationships among French physicist Henri Becquerel discovered
organisms. radioactivity in 1896, and, at the turn of the cen-
tury, Marie and Pierre Curie successfully sepa-
rated several radioactive elements. The idea of
dating rocks by using the rate of accumulation of
See Also helium produced by the decay of uranium was
conceived by English physicist Sir Ernest Ruth-
▶ Genotype erford in 1904. In the early part of the twentieth
▶ Phenotype century, J.J. Thompson, A.J. Dempster, and
▶ Phylogeny F.W. Aston developed the first mass spectrome-
▶ Species ters. Subsequently A.O. Nier built an instrument
948 Geochronology
that could be used to determine precise isotopic of any other loss or gain (the closed-system
ratios. He is also credited for the determination of assumption), the proportion of parent atoms
the isotope compositions of most elements in the P (or radioactive nuclides) disappearing per unit
1930s and 1940s. Between 1920 and 1953, sig- of time t is constant:
nificant advances in the understanding of the
potential of Pb isotopes for chronology are due dP
¼ l (1)
to A. Holmes, H.N. Russell, F.G. Houtermans, Pdt
and E. Gerling. In 1955, C.C. Patterson obtained
the first Pb-Pb ▶ isochron age on meteorites and The number of decay events per unit time dP/
dated the Solar System at 4.55 Ga, a number that dt is known as the “activity” of the radioactive
has hardly changed today. G.W. Wetherill should nuclide. For a number of parent atoms P = P0 at
be credited for the introduction of the Concordia time t = 0, this differential equation is equivalent
in 1956 and, together with T. Aldrich, for to
pioneering the 40 K-40Ar and the 87Rb-87Sr
methods. The merit of discovering extinct radio- P ¼ P0 elt (2)
activities falls on J.H. Reynolds in 1960 for
129 129 To determine the age of a system from the
I- Xe and on T. Lee and
D.A. Papanastassiou in 1974 for 26Al-26 Mg. In measurement of the number of parent atoms at
1961, L. Nicolaysen proposed the first isochron the present time, we must know P0, and therefore,
for the 87Rb-87Sr pair. R. Castaing and in this form, this equation is not a chronometer
G. Slodzian constructed the first effective ion (a notable exception is the 14C method). For each
probe in the early 1960s and in the late 1970s, parent atom, a daughter atom (or radiogenic
which opened the way to the in situ U-Pb ages of nuclide) is created, usually of a single element,
zircons, developed then by W. Compston. whose number can be denoted D. In a closed
C. Merrihue and G. Turner invented the system and for a stable daughter nuclide D, the
39
Ar-40Ar method in 1965. In 1969, number of parent and daughter atoms is constant;
D.A. Papanastassiou and G.W. Wasserburg hence,
constructed the first computer-assisted mass
spectrometer, which established the post-Apollo
D ¼ D0 þ P0 P ¼ D0 þ P elt 1 (3)
standard for isotope geochemistry. The first
ICP-source mass spectrometers dedicated to iso-
topic measurements and geochronology appeared The term P(elt 1) is a measure of the accu-
in the mid-1990s. mulation of the radiogenic nuclide during time t,
and therefore,

Overview 1 D D0
t ¼ ln 1 þ (4)
l P
Radioactive decay is a random process that
reflects the instability of some nuclei. Its proba- Even if D and P are measured, this equation is
bility of occurrence per unit of time is indepen- no more a timing device than Eq. 2 unless we
dent of time: atoms do not age. This probability, know the number of daughter atoms D0 at time
also known as the decay constant l, is specific to t = 0.
each radioactive nuclide. A parameter related
to the decay constant is the ▶ half-life T1/2,
which is equal to ln 2/l. Radioactive decay is “Rich” Chronometers: U-Pb and K-Ar
described as a random “Poisson” process, in
which the number of events is proportional to A first class of chronometers arises when the
the duration of the observation. In the absence condition D 0 D or
Geochronology 949

D P elt 1 (5) 206
Pb
y¼ 238 U
¼ el238 U T 1 (7b)
206 238
applies, as for the Pb- U dating of zircons, a
ubiquitous mineral of granites with formula where T stands for the geological age and the
ZrSiO4 into which U4+ easily substitutes but not asterisk refers to radiogenic Pb (total Pb minus
Pb2+. With this condition applying, Eq. 4 common or contamination Pb). The main limita-
becomes tion of this method is the presence of common Pb
introduced by inclusions, impurities, or contami-
206
nation during mineral processing.
1 Pb
t¼ ln 1 þ 238 (6) The 40K-40Ar method also belongs to this fam-
l238 U U
ily and dates the isolation of the host mineral.
These days, it is mostly used in its 39Ar-40Ar
When y = 206Pb/238U and x = 207Pb/235U are version, i.e., after irradiation of the sample in a
plotted against each other, the widely used nuclear reactor, fast neutrons with an energy
G
concordia diagram is obtained (Fig. 1). The >1 MeV react with 39K, which is in constant pro-
concordia is the locus of points for which the portions with 40K, to produce 39Ar. The age is
two ages of zircons agree, i.e., they are deduced from the 40Ar/39Ar ratio, in lieu of
concordant: 40
Ar/40 K, after proper calibration of the yield
of the nuclear reaction through a mineral standard
207
Pb of known 40K-40Ar age Tm referred to as the
x¼ 235 U
¼ el238 U T 1 (7a)
monitor:
1.2
Jack Hills zircons, Australia 4.5

1.0 Concordia
4.0
0.8
3.5
206Pb/ 238U
3.0
0.6
2.5
0.4 2.0
1.5
0.2 1.0
0.0
0 10 20 30 40 50 60 70 80 90
207Pb/ 235U
Geochronology, Fig. 1 The concordia plot, with U and zircons plot significantly off the concordia, the age used
Pb analyses of two detrital zircon grains from the Jack in practice is deduced from the upper intercept with the
Hills (Western Australia) by Wilde et al. (2001). The line going through the origin (207Pb*/206Pb* age, dashed
concordia is the locus of points for which the 206Pb-238U lines)
and 207Pb-235U ages (here shown in Ga) agree. When
950 Geochronology
ð40 Ar=39 ArÞspl el40 K T spl 1 easily, because it is a rare gas with no ionic or
¼ (8) covalent bonding in crystals: only the weak van
ð40 Ar=39 ArÞ m el40 K T m 1
der Waals forces bind Ar to other species.
A major advantage of this technique is that
resetting of the chronometer by thermal reheating
or weathering and the presence of excess Ar can “Poor” Chronometers: Isochrons
be easily identified. Argon is released by laser
ablation or, most commonly, by stepwise heating: For a second class of chronometers, the condition
undisturbed minerals show very large “plateaus” D0 D does not apply, and we replace it by the
with 40Ar/39Ar remaining constant over a broad principle of isotopic homogenization. Isotope
range of 39Ar degassing, whereas, for perturbed fractionation (both natural and analytical) is sim-
samples, 40Ar/39Ar reflects old ages only for ply corrected for by internal normalization
high-temperature steps. In the case of inherited against some arbitrary reference ratio of stable
Ar, the high-temperature steps may be anoma- isotopes, e.g., 179Hf/177Hf = 0.7325 for Hf. In
lous. Figure 2 shows a 39Ar-40Ar dating of the marine carbonates, the 87Sr/86Sr ratio is exactly
famous Martian meteorite ALH84001 (Bogard the same in calcite as in the seawater from which
and Garrison 1999), which has been suggested it precipitated; as the mantle melts, 176Hf/177Hf is
to harbor microfossils. A limitation of this tech- the same in the molten liquid as in the residue. Let
nique is the presence of atmospheric 40Ar, which us therefore divide Eq. 3 by the number D0 of
is corrected for by measuring 36Ar and using atoms of a stable isotope of the same element as
(40Ar/40Ar)atm = 295.5. Argon loss also happens the radioactive nuclide represented by D.
5.5 39
Ar-40Ar plateau age of ALH 84001
5.0
4.3 Ga
Age Ga
4.5
4.0
3.5
0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1

Cumulative fraction of 39Ar outgassed
Geochronology, Fig. 2 39Ar-40Ar in the gas fractions are plotted as a function of the progressive release of 39Ar.
released by stepwise heating of the Martian meteorite A plateau age (here 4.3 Ga) indicates that large fractions
ALH 84001 (Bogard and Garrison 1999), which has of the sample kept the memory of the same chronological
been suggested to host microfossils. By irradiation in a event. Anomalous ages at low temperatures (small frac-
nuclear reactor, fast neutrons change the 39K of the sample tions of 39Ar released) indicate partial resetting by
(which is in constant proportion with 40K) into 39Ar. The weathering or reheating. Anomalous ages at high temper-
39
Ar/40Ar is a substitute for the 40K/40Ar ratio. The ages atures indicate trapping of anomalous Ar components
Geochronology 951
Because the system is closed, D0 remains con-

and a similar isochron based on the system
stant, and therefore, 147
Sm-143Nd date 2.1 Ga old Birimian basalts
from West Africa (Blichert-Toft et al. 1999).
D D P lt
0 ¼ 0 þ 0 e 1 (9) D/D0 represents the ratio of the radiogenic
D t D 0 D t nuclide to its stable isotope (e.g., 176Hf/177Hf),
and P/D0 is the “parent/daughter” ratio, in most
This is the standard “isochron” equation. cases proportional to an elemental ratio (e.g.,
Figure 3 shows how an isochron based on the 176
Lu/177Hf). If two samples 1 and 2 formed at
system 176Lu-176Hf: the same time from an isotopically homogeneous
176 176 176 medium (ocean, magma), they share the same
Hf Hf Lu l176 t
(D/D0 )0, and, taking the 176Lu-176Hf system as
177 Hf
¼ 177 Hf
þ 177 Hf
e Lu 1
t 0 t an example, the time is obtained from
(10)
G
0.2845
Birimian basalts Ga
0.2840 .15
(West Africa) 2
T=
0.2835
176Hf/177Hf
0.2830
0.2825
0.2820
0.2815
0.00 0.01 0.02 0.03 0.04 0.05 0.06 0.07
176Lu/177Hf
0.5150
0.5140
143Nd/144Nd
0.5130
0.5120
Ga
2.10
T=
0.5110
0.5100
0.00 0.05 0.10 0.15 0.20 0.25 0.30 0.35
147Sm/144Nd
Geochronology, Fig. 3 Two examples of isochrons, Africa (Blichert-Toft et al. 1999). Slopes of these iso-
using the 176Lu-176Hf (top) and 147Sm-143Nd (bottom) chrons give the age at which these volcanic rocks were
systems on the same early Proterozoic basalts from West extracted from the mantle
952 Geochronology
" 176 #
1 Hf=177 Hf 2 176 Hf=177 Hf 1 The asterisk stands for radiogenic Pb. Here the
t¼ ln 1 þ 176 two parent nuclides on the one hand and the two
l176 Lu ð Lu= Hf Þ2 ð Lu= Hf Þ1
177 176 177
daughter nuclides on the other hand are isotopes
(11) of the same element, which dispenses the analyst
from measuring the U and Pb contents. In a plot
This age dates the time at which the two sam-
of 207Pb/204Pb versus 206Pb/204Pb, samples
ples last shared the same 176Hf/177Hf ratio, and
formed at the same time from the same reservoir
their parent/daughter ratios (here Lu/Hf) were
define a straight line (another isochron) (Fig. 4),
fractionated. This method is commonly used for
and the formation age can be retrieved from the
parent/daughter systems with a long half-life,
slope. Historically, this is how Clair C. Patterson
typically 147Sm-143Nd, 176Lu-176Hf, and
187 determined the age of the Solar System. For sta-
Re-187Os. The 87Rb-87Sr technique is broadly
tistical reasons, it is today common practice to
used to date medium-temperature metamorphic
use a 207Pb/206Pb versus 204Pb/206Pb isochron
events, 147Sm-143Nd for the emplacement of
plot instead, in which the intercept, and not the
ancient basalts and komatiites, 176Lu-176Hf in
slope, of the isochron gives the 207Pb*/206Pb*
garnets for high-temperature (>600 C) meta-
ratio and therefore the age. Figure 4 shows an
morphic events, and 187Re-187Os for the deposi-
isochron representing the Pb-Pb data on chon-
tion of reduced sediments (black shales) and
drules, little spherical blebs of molten material,
some ores.
and calcium-aluminum-rich refractory inclusions
A particular application is the Pb-Pb method,
(▶ CAIs) from the ▶ carbonaceous chondrite
which combines the two chronometers
206 Allende (Connelly et al. 2008), which represents
Pb-238U and 207Pb-235U. Equation 6 for the
one of the finest attempts to date an early sample
former system slightly modified as
of planetary material.
207 207
Pb Pb
204
204
Pb t Pb 0 Extinct Radioactivities
235
U
el235U 1
t
¼ 204 (12) A third class of chronometers relevant to
Pb t
astrobiology is that of extinct radioactivities.
is divided by the equation for the latter These have a short half-life (and therefore a
large l). For large values of lt, P becomes negli-
206 206 gible, and therefore the closed-system condition
Pb Pb
204
204 reads
Pb t Pb 0
238
U Dtoday ¼ Pt þ Dt (15)
el238U 1
t
¼ 204 (13)
Pb t
Let us write this equation for a sample (spl)
to give and divide it by the abundance (D0 )t = (D0 )today
of a stable isotope of D
207
Pb=204 Pb t 207 Pb=204 Pb 0 207 Pb spl Earth Earth 0 spl
206 ¼ D D P P
Pb=204 Pb t 206 Pb=204 Pb 0 206 Pb ¼ þ
1 el235 Ut 1 D0 today D0 t P0 t D0 today
¼ l
137:88 e 238 Ut 1 (16)
(14)

in which we introduce P0 , astable isotope of the
where we used the observation that, today, the parent nuclide Pt ¼ P0today . Using (Eq. 9), this
0
238
U/235U ratio is constant and equal to 137.88. equation is equivalent to
Geochronology 953
0.90 The age of Allende carbonaceous chondrite

Chondrules
Refractory inclusion (CAI)
0.85
207Pb/ 206Pb
0.80
0.75
0.70
0.65
4,567 Ma
G
0 0.01 0.02 0.03 0.04 0.05 0.06
204Pb/ 206Pb
Geochronology, Fig. 4 A Pb-Pb “inverse” isochron of the carbonaceous chondrite Allende. The age given by the
refractory calcium-aluminum-rich inclusions and chon- intercept (no stable 204Pb, so Pb is purely radiogenic)
drules (small blebs of molten silicates) (Connelly gives 4,567 Ma, one of the oldest estimates of the age of
et al. 2008), which accounts for most of the material in the Solar System
spl Earth Earth spl Earth

D D P ð182 W=183 WÞtoday ð182 W=183 WÞtoday
¼ þ
D0 today D0 today P0 t ð180 Hf=183 WÞspl 180 Hf=183 WÞEarth
today ð (19)
" 0 Earth # 182 spl ¼ Earth 182 Earth
today
P0
spl
P ¼ 180 Hf ¼ 180 Hf el182Hf t
(17)
D0 today D0 today
Hf t Hf 0
Extracting time from this equation

For the example of the 182Hf-182 W chronom- requires that (182Hf/180Hf)0 of the Earth
eter (T 1/2 = 8.9 Ma) (Fig. 5), this equation reads is known. However, dividing this equation
182 spl 182 Earth 182 Earth for the core (182Hf 0) by the same equation
183
W
¼ 183
W
þ 180
Hf for the mantle gives the age mantle-core
W today W today Hf t
separation.
spl 180 Earth (18)

180
Hf

Hf This method is employed for a number of
183 183
W today W today “extinct” short-lived nuclides. The chronometers
26
Al-26Mg (T1/2 = 700,000 a), 60Fe-60Ni
This is the equation of the extinct radioactivity (T1/2 = 2.6 Ma), 53Mn-53Cr (T1/2 = 3.7 Ma),
isochron. The conventional isochron cannot be 182
Hf-182 W (T1/2 = 8.9 Ma), and 129I-129Xe
used when all the parent nuclides have decayed (T1/2 = 15.7 Ma) provide essential chronological
away. When 182 W/183 W is plotted against information on the accretion of planetary mate-
180
Hf/183 W (note that the two nuclides of the rial from the solar nebula, its thermal history, and
latter ratio are both stable), samples formed at metal-silicate (core-mantle) segregation, while
the same time in isotopic equilibrium within the the 146Sm-142Nd (T1/2 = 103 Ma) system proved
Earth (e.g., mantle and core) define a straight line. essential to demonstrate the existence of very
The slope of this isochron varies with time as early crust.
954 Geochronology
1.8525
n
ro
ch
iso
1.8520
eit
dr
on
Ch
182W/183W
1.8515 30 Ma Bulk silicate

earth
1.8510
1.8505
0 Solar 10 20 30 40
nebula
180Hf/183W
Geochronology, Fig. 5 Dating of core-mantle segrega- into the silicate. The point of intersection of the two lines
tion by the 182Hf-182 W method (Yin et al. 2002). The represents the unfractionated solar nebula. The difference
difference in slope between the alignment of the points in slopes show that core-mantle separation in the Earth
representing the chondrites and that representing the sili- occurred some 30 Ma after the formation of the Solar
cate Earth is a measure of the last fractionation of the System
Hf/W ratio. Tungsten fractionates into the metal and Hf
Closure Temperatures If cooling is fast, the radiometric system dates the

time at which temperature in the rock dropped
What chronology dates is actually not discrete below a particular value. The transition is usually
formation events. Upon cooling, minerals fast (a few tens of degrees). Typical closure tem-
keep exchanging radiogenic isotopes with peratures vary from 1,000 C for U-Pb in zircons
their surroundings: the chronometric system is to 800 C for Pb-Pb in pyroxene and Lu-Hf in
open. In most cases, what a chronometer garnet and to 300 C for K-Ar in feldspar and
records is therefore the time at which Re-Os in sulfides. If cooling is very slow, the
diffusion stopped (Fig. 6). Diffusion of a system remains ajar for a long time, the demise
particular ion is a thermally activated molecular of diffusion is sluggish, and the significance of
process of exchange between adjacent minerals the date is blurred.
or between a phase and the interstitial Diffusion loss is one of the major issues of
medium and is characterized by the energy of geochronology. Rocks from the sub-lithospheric
activation E and the so-called pre-exponential mantle evolve for long times above the
term D0. If cooling is fast, the closure temperature closure temperature and therefore cannot be
Tc (in K) is given by Dodson’s (1973) implicit dated by isotopic chronometers. The closure
equation: temperature varies as 2 ln a, so chronometers in
smaller systems, first and foremost isolated
E crystals, close at a lower temperature than in
Tc ¼
(20)
R ln AD0 RT c =Ea2 ðdT=dtÞ
2 the larger ones and may be easily reset by
mild thermal events. Isochrons of large whole-
in which R is the gas constant, a the grain size, d rock samples are often used to mitigate this
T/dt the cooling rate, and A a constant equal to 55. problem.
Geochronology 955
Geochronology,
Fig. 6 Cooling age and Open system Closed system
closure temperature of a
chronometric system with
Amount of radiogenic
decay constant l. The
nuclide remaining
cooling time and the
in the mineral
corresponding cooling age
tc are defined by linearly
extrapolating the ingrowth
of radiogenic nuclides at Temperature T
zero, while the closure λP
temperature Tc is the
temperature of the mineral Closure temperature
Tc
at this time. The system is
open and looses the
daughter isotope for T > Tc
and is closed for T < Tc. G
The slope of the daughter
isotope evolution curve is
the activity lP Cooling age, tc
Time, t
Basic Methodology coil submitted to rapidly fluctuating currents to

ionize the other elements: this relatively new
Choosing a particular chronological system to technique accounts for most measurements for
use for a given problem depends on the duration Hf, W, Mg, and Fe. Sputtering of the sample by
of the age or age difference to be assessed. a 30 mm primary ion beam (SIMS or ion probe)
Enough decay must have happened that it mea- is the method of choice for the in situ measure-
surably affects the abundance of the daughter ment of U-Pb ages in zircons (Fig. 8).
isotope. In practice, a chronometer gives no use- The precision of measurements is first and
ful information if this interval exceeds five to foremost limited by counting statistics: a mass
eight half-lives. spectrometer can be seen as a device counting
The elements for which the isotope composi- ions arriving on a detector. The measurement
tion is sought are isolated in a variety of ways. can therefore be seen as another Poisson process,
Gases are extracted by heating and purified in which for n “events” must be associated with a
pffiffiffi
high vacuum enclosures. Solid samples are nor- standard deviation of n and a relative error of
pffiffiffi
mally digested with strong acids (HF, HNO3, 1= n . Let us assume that we use an ion probe
HCl) in pressurized vessels and the metals usu- with a 30 mm beam size to measure the abundance
ally separated by ion chromatography. Isotope of 207Pb in a zircon crystal with density of 4.65 g
ratios are measured on mass spectrometers, cm3 and which contains 100 ppm Pb. We also
which combine a source of ions, electrostatic assume that the beam excites a hemispherical
acceleration, a magnetic filter, and a set of detec- volume below the spot and a 1 % ion yield. We
tors. These instruments differ by the nature of must therefore divide the weight of sputtered
their ion sources. Gas sources use the bombard- material by the molecular weight of Pb and
ment of an electron beam to ionize atomic gases multiply by the isotopic abundance of 207Pb and
and are used for O, C, He, Ar, Xe, etc. Thermal finally multiply by the Avogadro number
3
ionization mass spectrometers (TIMS) produce to obtain 0:01 ð2=3 3:14Þ 15 104
ions by evaporative ionization at the surface of a 4:65 100 106 =207:2 0:22 6:0 1024 ¼ 2:1 108
hot filament and are used for Sr, Nd, Pb, and counts (approximate numbers suffice), i.e., an
Os. Inductively coupled plasma mass spectrome- excellent precision of 0.007 %. Reducing the
ters (ICP-MS) (Fig. 7) use Ar ions produced in a beam size to 1 mm would deteriorate the precision
956 Geochronology
Geochronology,
Fig. 7 The Nu 1,700 large
radius MC-ICP-MS
(multiple-collector
inductively coupled plasma
mass spectrometer) of
Ecole Normale Supérieure
in Lyon, France. The
essential parts of the mass
spectrometer are shown.
This instrument can
measure the isotope
composition of most ions
on quantities as small as
10 ng (one billionth of a
gram) with a precision of
0.005 % or better
Geochronology,
Fig. 8 The Cameca 1280
ion probe (secondary ion
mass spectrometer) of the
Centre de Recherches
Pétrographiques et
Géochimiques de Nancy,
France
to 1 % and make the measurement essentially contamination and common Pb can be removed
worthless. The quest for ages on smaller and and hence constitutes a significant source of
smaller objects is intrinsically limited by the uncertainty on ages. A third problem affects the
desired precision. chronometers based on beta decay: the kinetic
A second stringent limitation is contamina- energy of some emerging electrons is so small
tion, which for many elements such as Pb forces that they go undetected, and the corresponding
the analyst to work in ultra-clean environments decay constant cannot be accurately determined.
with filtered air and distilled reagents. Not all This is a limitation for the 87Rb-87Sr,
Geological Time Scale, History of 957
176
Lu-176Hf, and 187Re-187Os techniques, which Dodson MH (1973) Closure temperature in cooling geo-
can be mitigated to some extent by a cross- chronological and petrological systems. Contrib Min-
eral Petrol 40:259–274
calibration with U-Pb ages. Faure G, Mensing TM (2004) Isotopes. Principles and
Finally, a general limitation arises when data applications. Wiley, New York, p 897
expected to form an alignment actually scatter Ludwig K (2003) ISOPLOT: a geochronological toolkit
beyond analytical errors. This can be due to for Microsoft Excel 3.00. In Berkeley Geochronology
Center Special Publication No. 4, 2455 Ridge Road,
(1) a lack of isotopic homogeneity at t = 0; Berkeley 94709
(2) to the failure of the closed-system assumption Wilde SA, Valley JW, Peck WH, Graham CM (2001) Evi-
because of weathering or thermal resetting; and dence from detritical zircons for the existence of con-
(3) to a long duration of the event to be dated, e.g., tinental crust and oceans on the Earth 4.4 Gyr ago.
Nature 409:175–178
in case of slow cooling. The literature on least- Yin Q, Jacobsen SB, Yamashita K, Blichert-Toft J,
square regression with correlated or Télouk P, Albarède F (2002) A short timescale for
noncorrelated errors is abundant, but a standard terrestrial planet formation from Hf-W chronometry
reference is that of York (1969). Among all the of meteorites. Nature 418:949–952 G
York D (1969) Least squares fitting of a straight
codes used to evaluate alignments, the most line with correlated errors. Earth Planet Sci Lett
broadly used software is ISOPLOT created by 5:320–324
Ludwig (2003).
See Also Geological Time Scale, History of
▶ Earth, Age of Pierre Savaton

▶ Isochron Université de Caen Basse-Normandie, Caen,
▶ Isotope France
▶ Isotopic Ratio
▶ Mass Spectrometry
▶ Radioactivity Keywords
Chronology; Radiometric dating; Stratigraphic

scale
Albarède F (2009) Geochemistry: an introduction. Cam-

bridge University Press, Cambridge, p 356
History
Allegre CJ (2008) Isotope geology. Cambridge University
Press, Cambridge, p 512 Geology is the first historical science. Concept of
Blichert-Toft J, Albarède F, Rosing M, Frei R, Bridgwater time is a key concept in geology. Even if some
D (1999) The Nd and Hf isotopic evolution of the
geological processes operate over time of a few
mantle through the Archean. Results from the Isua
supracrustals, West Greenland, and from the Birimian seconds (e.g., earthquake waves), most of them
terranes of West Africa. Geochim Cosmochim Acta operate over million to hundreds of millions of
63:3901–3914 years. Thus, to tell the Earth’s history, we need to
Bogard DD, Garrison DH (1999) Argon-39-argon-40
build a geological time scale to place geological
“ages” and trapped argon in martian shergottites,
chassigny, and Allan hills 84001. Meteorit Planet Sci events and measure their length. This scale
34:451–473 results from a combination of time and space
Connelly JN, Amelin Y, Krot AN, Bizzarro M (2008) relations between rock formations (relative age)
Chronology of the solar system’s oldest solids.
and from the measure of spontaneous decay of
Dickin AP (1999) Radiogenic isotope geology. Cam- radioactive nuclides contained in rocks (absolute
bridge University Press, Cambridge, p 490 age). The only record we have of past geological
958 Geological Time Scale, History of
events is the rock preserved from erosion, alter- might be considered a precursor of the idea of
ation, and plate tectonics. index fossil, that is, fossils of organisms that lived
The geological time scale was first a chrono- in a short time span but widely distributed geo-
logical one, without date. Study of sedimentary graphically. But the time-keeper aspect of fossil’s
rocks had allowed first “geologists” to establish records remained marginal for a long time.
the simple basis for the stratigraphic scale. German geologist and mineralogist Abraham
Danish naturalist Nicolas Steno (1638–1686) G. Werner still had neglected them in his Kurze
noted that gravels, sands, and clays were laid Klassifikation und Beschreibung der
down in more or less horizontal layers, which he verschiedenen Geb€ urgsarten (1783), a strati-
called strata. In 1669, in his Prodromus (De graphical framework attempt that was widely
solido intra solidum naturaliter content adopted like scheme of the succession of geolog-
dissertationis prodromus), he enunciated princi- ical times. Fossils were just used to characterize
ples of stratigraphy: original horizontality, super- his formations, but no more. Faunal succession,
position, and lateral original continuity of layers. as the lithologic succession, could be used
Using these principles, it was possible to arrange directly to identify rocky formations. This
layers in a chronological succession, and some approach was that used by British geologist Wil-
attempts were tempted and offered local descrip- liam Smith to build his Geological Map of
tions of the pile of stratas. Steno identified in England and Whales and part of Scotland
these successions some angular unconformities, (1815). His Strata identified by organized fossils
which represented breaks in the stratigraphical (1816–1819) paved the way for the development
record, but without understanding their possible of the biostratigraphy. George Cuvier’s and
use in dating of geological events, even if he Alexandre Brongniart’s map and Description
showed that “geology” revealed a history. By géologique des environs de Paris (1811) proposed
contrast, he recognized that fossils were the to use fossils to distinguish marine from conti-
remains of once living organisms. In 1667, work- nental sediments and to explain succession of
ing in Tuscany, he noted that shark teeth bored a sedimentological events. They used them as indi-
striking resemblance to some stony objects found cators of paleoecological conditions rather than
embedded within rock formations, called “tongue strict stratigraphical tools.
stones.” In a few decades in the early 1800s, the geol-
English philosopher Robert Hooke ogists who mapped the world established a time
(1635–1703) considered fossil’s shells as medals scale based on assemblages of fossils, far more
or monuments of nature, as greatest and more complex than the earlier succession of rocks
lasting monuments and records of antiquity. He (primary, secondary, tertiary) of the 1700s
claimed that animals living today might not have based on a lithological time scale.
been alive in the past and animals presented in the Time and history of the Earth was divided in
past were no longer present today. He interpreted time units chronologically distinguished both by
the differences between past and actual faunas by their vertical spatial relationships and their fossils
imagining a kind of transformation of species in content. Fossils determined a major division
time. This idea was taken back later by Leibniz between the early time of the Earth, without vis-
before being developed and theorized by Jean- ible fossils and the Phanerozoic time (today the
Baptiste Lamarck. But it was too premature for fourth eon) characterized by its fossil diversity.
the 1600s. Hooke used for the first time the word Phanerozoic was subdivided into eras, periods,
“chronology” and suggested a powerful tool to and epochs. The development of a systematic
date geological layers. These medals (fossils) palaeontology during the nineteenth century and
allowed establishing continuity between layers studies of some of the less visible one revealed
separated by great distance or layers of different new fossil discontinuities in the time and justified
lithology. They allowed replacing geographical new subdivisions. French naturalist Alcide
isolated layers in a succession. Robert Hooke d’Orbigny showed how microscopic fossils,
Geological Time Scale, History of 959
especially Foraminifera, could be used to charac- time had been molten. Solid today, still hot, it
terize Cenozoic stratas. Paleontologists who stud- would lose heat by conduction, compatible with
ied the Jurassic ammonites identified species with the increasing temperatures recorded in mines or
a very short time range and suggested to use them boreholes. In 1862, using Fourier’s laws of
to define small biostratigraphic units (biozones). cooling, Thomson estimated the Earth’s center
The theory of the Evolution of Darwin temperature to about 3,800 C and the age of
(1859) wasn’t necessary to think a biostrati- the Earth to about 20–400 million years. In
graphic scale, but gave a new breath to the fossil’s 1897, Thomson, now Lord Kelvin, ultimately
tool. The succession of changing fauna could be settled on an estimate that the Earth was 20–40
explained by successive destructions and crea- million years old.
tions, migrations, or evolution. In the continuity The discovery of radioactivity by Henri Bec-
of Cuvier thinking, most of the first stratigraphers querel in 1896 and the isolation of radium by
favored extinctions. Proposing that new species Pierre and Marie Curie in 1898 changed the clas-
emerged as a result of modifications to earlier sic view of a cooling Earth and offered to geology G
species was more powerful. But the theory of the tool it needed to measure time. The work of
evolution obliged to reconsider the age of the Rutherford and Soddy about radioactive decay
Earth, because progressive changes required series and the discovery that lead was the end
longer time. product of the uranium decay chain by American
The Archbishop Ussher of Ireland wrote in chemist Bertram Boltwood in 1905 stated the
1664 that the Earth was created 4,004 years fundamental basis of radiometric dating and
before Christ. He had not been alone to interpret allowed geologists to put absolute dates in their
the Bible word for word to determine the antiq- geological column. Using what he knew about
uity of the Earth. All these biblical ages were very radium-lead decay, Boltwood aged rocks from
short. George Buffon, who believed in a cyclical 250 million to 1.3 billion years old. Although
history of the Earth, clearly longer than that given his estimate of the Earth’s age was inaccurate,
in the biblical accounts, published in his Epoques his paper, published in 1907, made clear that
de la Nature (1778) an empirical calculation of sediments from the same layers had similar
the age the Earth. Following the idea that the uranium-lead ratios and that generally samples
Earth might originally have been molten, he mea- from younger layers had a lower proportion
sured the cooling time of iron balls of different of lead.
sizes, and other materials, and concluded through In 1911, British geologist Arthur Holmes
this model that Earth should have been determined an age of 1,640 million years for
75,000 years old. This duration was to great Archean rocks and published in 1913, in The
enough for many of his contemporaries. Age of the Earth, the first stratigraphic time
The uniformitarian’s ideas spread by British scale with radioactive ages for the boundaries of
geologist Charles Lyell suggested estimating the the main units. He estimated the beginning of the
actual age of the Earth using sediment accumu- Phanerozoic time at about 600 million years
lation or denudation rate as indicators of time. before present (today this limit is fixed at
John Phillips transformed the unit of geological 542 Ma). Arthur Holmes’s time scale established
time into an equivalent term of years and pro- in 1947 was very similar than these of today.
posed in 1860 in Life on the Earth an age of Working on lead isotopes in iron and stone mete-
Phanerozoic time between 38 and 96 million of orites, American geochemist Clair Patterson
years. It wasn’t enough for explaining the established and published in 1956 an age of the
evolution theories of Darwin but compatible Earth of 4,550 70 million years.
with ages calculated by British physicist William With fossil’s chronology of sedimentary rocks
Thomson (better known as Lord Kelvin) through and radioactive age of crosscutting igneous
his model of a cooling Earth. Most of the geolo- rocks, geologists have built the geological time
gists of the 1800s believed that Earth of the first scale. Earth history could finally be told.
960 Geological Timescale
See Also timescale. His book, The Age of the Earth (1913,
2nd edition 1937), written when he was only
▶ Cuvier’s Conception of Origins of Life 22, had a major impact on those interested in
▶ Darwin’s Conception of the Origins of Life ▶ geochronology. For his pioneering scale,
▶ Earth, Age of Holmes carefully plotted four radiometric dates,
▶ Geochronology one in the Eocene and three in the Paleozoic
▶ Geological Timescale from radiogenic helium and lead in uranium min-
▶ Lamarck’s Conception of Origins of Life erals against estimates of the accumulated
maximum thickness of Phanerozoic sediments. If
we ignore sizable error margins, the base of Cam-
brian interpolates at 600 Ma, curiously close to
Brush S (1996) Transmuted past: the age of the Earth and modern estimates (541 1.0 Ma). The new
the evolution of the elements from Lyell to Patterson. approach was a major improvement over a
Cambridge University Press, New York previous “hour-glass” method that tried to esti-
Jackson PW (2006) The chronologers’quest. The search mate maximum thickness of strata per period to
for the age of the Earth. Cambridge University Press,
New York determine their relative duration but had no way of
Lewis C, Knell S (eds) (2001) The age of the Earth: from estimating rates of sedimentation independently.
4004 BC to AD 2002. Geological Society Special
Publication, London
Oldroyd D (2006) Earth cycles, a historical perspective.
Greenwood Press, London Overview
Rudwick M (2008) Worlds before Adam. The reconstruc-
tion of geohistory in the age of reform. University of Calibration to linear time of the succession of
Chicago Press, Chicago events recorded in the rocks on Earth has three
components: (a) the international stratigraphic divi-
sions and their correlation in the global rock record;
Geological Timescale (b) geochronology, the means of measuring linear
time or elapsed durations from the rock record; and
Felix M. Gradstein (c) the methods of effectively joining the two
University of Oslo, Blindem, Oslo, Norway scales, the stratigraphic one and the linear one.
For clarity and precision in international com-
munication, the rock record of Earth history is
Definition subdivided into a “chronostratigraphic” scale
(Fig. 1) of standardized global stratigraphic units,
The Geologic Time Scale is the framework for such as “Devonian,” “Miocene,” “Zigzagiceras
deciphering and understanding the long and com- zigzag ammonite zone,” or “polarity Chron
plex history of the Earth. Understanding the C25r.” This chronostratigraphic calendar is not
physical, chemical, and biological processes unlike a historical calendar in which civilization
since the Earth formed requires a detailed and periods, such as the Minoan Period, Reign of Louis
accurate timescale. The timescale is the tool XIV, or American Civil War are used as building
“par excellence” of the geological trade blocks, devoid of a linear scale. Archeological
(Gradstein et al. 2004; Ogg et al. 2008). relics deposited during these intervals, such as the
Palace of Minos on Crete, Versailles, or spent
cannon balls at Gettysburg, comprise the associ-
History ated physical chronostratigraphic record.
The chronostratigraphic scale is thus assem-
British geologist Arthur Holmes (1890–1965) bled from rock sequences stacked and segmented
was the first to combine radiometric ages with in relative units based on their unique fossil and
geologic formations in order to create a geologic physical content. This scale becomes meaningful
Geological Timescale
961
Geological Timescale, Fig. 1

G
962 Geology of Mars
and useful through correlation of the unique fossil International Commission of Stratigraphy (ICS) geologi-
and physical record to other sections of sedimen- cal timescale at http://www.stratigraphy.org/index.
php/ics-chart-timescale
tary or volcanic rocks in outcrops or wells across Ogg JG, Ogg G, Gradstein FM (2008) The concise geo-
the globe. logic time scale. Cambridge University Press, Cam-
The standard chronostratigraphic scheme, in bridge, 177 p
downloadable graphic format and available from
the International Commission of Stratigraphy (ICS)
website (http://www.stratigraphy.org/index.php/ics- Geology of Mars
chart-timescale), is made up of successive stages in
the rock record, for example, Cenomanian, ▶ Areology
Turonian, then Coniacian, etc., within the Creta-
ceous System. The chronostratigraphic scale is an
agreed convention, whereas its calibration to linear Geomicrobiology
time is a matter of discovery or estimation. The
estimation comes from radiometric dating and Ricardo Amils
astrochronology (orbital tuning of long sets of Departamento de Biologia Molecular,
cycles) in the sedimentary rock record and is con- Universidad Autónoma de Madrid, Madrid, Spain
tinually refined. Thus, the chronostratigraphic chart
is updated, usually once a year.
In contrast to the Phanerozoic which has an Definition
agreed-upon chronostratigraphic scale with for-
mal stage boundary stratotypes, Precambrian Geomicrobiology examines the role of microbes in
stratigraphy is formally classified chronometri- geological processes. Examples of such processes
cally (see ▶ Proterozoic Eon, ▶ Early Archean). are the weathering of rocks, soil and sediment
formation and transformation, the genesis and deg-
radation of minerals, and the genesis and degrada-
See Also tion of fossil fuels. Due to its transdisciplinary
essence and the subject of study, geomicrobiology
▶ Archean Eon is of special astrobiological interest.
▶ Earth, Age of
▶ Geochronology
▶ Geological Time Scale, History of See Also
▶ Hadean
▶ Proterozoic Eon ▶ Anoxygenic Photosynthesis
▶ Bacteria
References and Further Reading ▶ Chemolithotroph
▶ Deep Subsurface Microbiology
Gradstein FM, Ogg JG, Smith AG, Agterberg FP, ▶ Early Archean
Bleeker W, Cooper RA, Davydov V, Gibbard P, ▶ Environment
Hinnov L, House MR, Lourens L, Luterbacher HP, ▶ Hot Spring Microbiology
McArthur J, Melchin MJ, Robb LJ, Shergold J,
Villeneuve M, Wardlaw BR, Ali J, Brinkhuis H, ▶ Iron
Hilgen FJ, Hooker J, Howarth RJ, Knoll AH, ▶ Iron Cycle
Laskar J, Monechi S, Powell J, Plumb KA, Raffi I, ▶ Lithotroph
Röhl U, Sadler P, Sanfilippo A, Schmitz B, Shackleton ▶ Magnetotactic Bacteria
NJ, Shields GH, Strauss H, Van Dam J, Van
Kolfschoten T, Veizer J, Wilson D (2004) A geologic ▶ Prokaryote
time scale 2004. Cambridge University Press, Cam- ▶ Sulfate Reducers
bridge, p 589, 200 figs and tables ▶ Sulfur Cycle
Geothermobarometers 963
from a hot interior. The Earth’s internal heat

Geotherm derives from a combination of residual heat from
planetary ▶ accretion (20 %) and heat produced
Daniele L. Pinti through radioactive decay of U, Th, and K (80 %).
GEOTOP Research Center for Geochemistry and The magnitude of the geothermal gradient
Geodynamics, Université du Québec à Montréal, depends on the rate of heat production at depth,
Montréal, QC, Canada the dynamics of the system, and the conductivity
of rocks. The highest gradients, 40–80 K km1,
are measured at oceanic spreading centers (▶ mid-
Definition ocean ridges) or at island arcs where magma is
close to the surface. The lowest gradients occur at
In the interior of a planet, the temperature ▶ subduction zones where cold lithosphere
increases with depth along a curve referred to as descends into the mantle. The gradient in old sta-
geotherm. It is the graphical representation of the ble ▶ continental crust is ~30 K km1 and is G
geothermal gradient. The shape of the curve somewhat lower in cratons. Upwelling parts of
depends upon the distribution of heat sources, the mantle ascend nearly adiabatically (i.e., they
the rate of heat production at depth, the dynamics lose little to no heat to the surroundings), and the
of the system (such as mantle convection), and gradient is very low, about 0.3 K km1.
the heterogeneities in the properties of rocks
(conductivity, compressibility).
See Also
See Also
▶ Continental Crust
▶ Heat Flow, Planetary
▶ Geothermal Gradient
▶ Mid-Ocean Ridges
▶ Subduction
Geothermal Flux
Geothermobarometers
Jean-Emmanuel Martelat
LST UMR5570, Université Claude Bernard
Lyon 1, St Martin d’Hères, Grenoble, France
Geothermal Gradient
Keywords
Nicholas Arndt
ISTerre, Université Grenoble Alpes, France Geobarometer; Geothermometer; Metamorphic
rocks; Metamorphism
Definition
Definition
A geothermal gradient is the increase in temper-
ature with increasing depth beneath the Earth’s Geothermobarometers refer to all types of reac-
surface. This gradient is due to outward heat flow tions that are useful to estimate temperature and
964 Geothermobarometers
pressure recorded in a magmatic or a metamor- Several chemical reactions can help constrain the
phic rock when it crystallizes or recrystallizes. temperature. One of the most broadly used
geothermometers is based on the diffusional
exchange of Fe2+ and Mg2+ between garnet and
Overview biotite. Fe2+ and Mg2+ cations are easily
exchangeable since they have similar electron
Minerals have different physical properties (e.g., valence and atomic radii. This exchange occurs
volume, density, or entropy). When pressure without change in size of both the garnet and the
increases, minerals such as quartz biotite. The reaction is
(density 2.66 g/cm3) may be metamorphosed
into high-pressure polymorphs (e.g., for quartz: Phlogopite þ almandine ¼ annite þ pyrope
coesite, density 3.0 g/cm3). This kind of reac-
tion is called polymorphic: the chemical compo- KMg3 AlSi3 O10 ðOHÞ2 þ Fe3 Al2 ðSiO4 Þ3
sition remains SiO2, but the atoms are rearranged ¼ KFe2þ
3 AlSi3 O10 ðOHÞ2 þ Mg3 Al2 ðSiO4 Þ3
in a more compact manner. On the other hand,
as temperature increases, minerals such as where phlogopite and annite are the Mg- and
kyanite (Al2SiO5) (standard entropy “S” at Fe-pure end-member of biotite whereas pyrope
T=298 83 J/mol-K) may be metamorphosed and almandine are the Mg- and Fe-pure
into a high-temperature polymorph, sillimanite end-members of garnet, respectively. When tem-
(Al2SiO5) (S at T=298 K 93 J/mol-K), perature decreases, the Fe/Mg ratio decreases in
displaying a higher atomic disorder. Hence, a biotite and increases in garnet. The most critical
mineral or a mineral assemblage is sensitive to assumption behind temperature and pressure
P-T conditions and is only stable within a given obtained by geothermobarometry is that equilib-
range of those conditions (see also “▶ metamor- rium reactions were frozen during cooling. As
phic rocks” and “▶ metamorphism”: metamor- diffusivity depends on temperature and on the
phic facies). Moreover, mineral compositions distance between the exchange zones in the bio-
corresponding to a specific chemical equilibrium tite and garnet in contact, one can obtain temper-
allow a quantitative estimate of P-T conditions: ature gradients from the rim to the core of the
this method is called geothermobarometry. We given mineral pairs. Other equilibriums can con-
use the term “geothermometer” for metamorphic strain temperature: e.g., polymorphic thermome-
reactions (or chemical equilibrium) that are highly ter, solvus thermometer (e.g., feldspar
sensitive to temperature and “geobarometer” for exsolution), net transfer reaction thermometer
reactions that are sensitive to pressure. (some minerals are consumed while others are
Our ability to estimate or calculate the quan- produced), trace element thermometer, and stable
titative P-T conditions of a given rock depends on isotope partitioning thermometer. Some other
(1) the observation of the sample’s texture under techniques, such as liquid–vapor homogenization
the microscope, (2) the quality of the experimen- points in fluid inclusions, provide quantitative
tal calibration of the chemical reactions (stability thermometry results. The best way to obtain
field of mineral phases), (3) the accuracy of the quantitatively the T and P recorded in rocks is
thermodynamic properties of the minerals to combine several internally consistent
(database), and (4) the precision of chemical ana- thermobarometers. Various freeware programs
lyses on the minerals (e.g., with electron micro- available on the Internet calculate numerous pos-
probes reaching a 1 mm2 spot resolution). On sible equilibriums with respect to an internally
well-studied samples bearing relevant mineral consistent thermodynamic database for a large
assemblages, quantitative results can be obtained number of minerals (e.g., TWEEQ, Berman
with an accuracy of 50 C and 1–1.5 kbars. 1991; THERMOCALC, Powell et al. 1998).
Geyser 965
See Also Definition
▶ Geothermal Gradient A geyser is a type of hot spring that intermittently

▶ Metamorphic Rock erupts hot water and steam. It is a temporarily geo-
▶ Metamorphism logical feature caused by hydrothermal activity in
▶ Plate Tectonics volcanic areas. The formation of geysers is related to
the uprising of pressurized water in rock conduits, at
superheated conditions. Decrease of pressure close
to the surface makes superheated water to rapidly
Berman RG (1991) Thermobarometry using change into steam (flashing) driving the hot water
multiequilibrium calculations: a new technique with eruption. After the pressure decreases to a threshold
petrologic applications. Can Mineral 29:833–855 value, the eruption stops and the system pressurizes
Powell R, Holland TJB, Worley B (1998) Calculating again to initiate the following eruption.
phase diagrams involving solid solutions via
non-linear equations, with examples using G
THERMOCALC. J Metamorph Geol 16:577–588
Spear FS (1993) Metamorphic phase equilibria Overview
and pressure-temperature-time paths.
Mineralogical Society of America Monograph, New
York, 799 pp The name geyser is derived from Geysir, a geyser in
southwestern Iceland which was the first geyser
described in written sources (1294 CE). The name
in Icelandic means “spurt out.” On Earth, there are
Germ two types of geysers: fountain geysers and cone
geysers. Fountain geysers erupt directly from a
▶ Microorganism pool of water, while cone geysers erupt from a
mound made of siliceous sinter (opal) called gey-
serite. Old Faithful, the most well-known geyser at
Yellowstone National Park, USA, is a cone geyser.
On Earth, geysers exist almost everywhere, where
German Aerospace Center there is a vigorous hydrothermal system, but a few
places concentrate large amounts of them. The larg-
▶ DLR, Germany est concentration of geysers in the world (300–500)
is at Yellowstone National Park USA, followed by
the Valley of Geysers in the Kamchatka Peninsula
of Russia (200 geysers) and El Tatio, Chile (80 gey-
Geyser sers), with many others in the Taupo Volcanic
complex, New Zealand, and in Iceland.
Daniele L. Pinti On Earth, geysers have an astrobiology inter-
GEOTOP Research Center for Geochemistry and est because, despite the harsh conditions, around
Geodynamics, Université du Québec à Montréal, geyser’s mounds and pools, thermophile
Montréal, QC, Canada (<70 C) and ▶ hyperthermophile (<110 C)
prokaryotes can develop. Thus, geysers are
niches favorable to extremophile life and likely
Keywords natural analogue of harsh conditions in the
Hadean and Archean Earth.
Hydrothermal environments; Extremophiles; There are other planetary bodies in the Solar
Cryovolcanism; Enceladus; Triton System showing jet eruptions similar to geysers.
966 Giant Impact
Icy-cold volcanism (▶ cryovolcanism) is also masses of at least a lunar mass and often signif-
able to reproduce geyser-like features called icantly larger. Planetary embryos are thought to
cryogeysers. The Voyager 2 flyby of Neptune in grow by runaway and ▶ oligarchic growth from a
1989 showed geyser-like eruptions on its moon swarm of small ▶ planetesimals primarily via
Triton. These eruptions are thought to be made of embryo-planetesimal collisions. Once the
liquid nitrogen and dust and they were driven by local density of embryos and planetesimals is
solar heating. Similar solar heating-driven eruptions comparable, embryos can grow by embryo-
are believed to occur in the south pole region of embryo collisions. These so-called giant impacts
Mars during spring thaw. Finally, pulsating bright are very energetic events whose consequences
plumes emanating from Enceladus’ south pole have are not completely understood. In fact, most
been discovered by the ▶ Cassini flyby in 2005. such impacts probably do not lead to perfect
These were interpreted as geysers drawing from a merging and probably generate significant
large body of liquid water. Tidal forces induced by amounts of collisional debris. The last giant
Saturn’s gravity probably create sufficient friction impact on Earth is thought to have been a
to partially melt the ice crust of Enceladus and to low-speed, off-center collision with a Mars-
create liquid water pools from which the geysers sized projectile, usually called Theia. The debris
originate. Thus, cryogeysers on icy moons could from this collision stayed in orbit around Earth
bring to their surface water and organics. and coalesced into the Moon.
See Also
▶ Cassini See Also

▶ Cryovolcanism
▶ Enceladus ▶ Impact, Hit and Run
▶ Hydrothermal Environments ▶ Moon, Origin of
▶ Hyperthermophile ▶ Oligarchic Growth
▶ Thermophile ▶ Planetary Embryo
▶ Triton ▶ Planetesimals
▶ Voyager, Spacecraft ▶ Runaway Growth
Kargel JS (1995) Cryovolcanism on icy satellites. Earth Giant Planets

Moon Planet 67:101–113
Therese Encrenaz
Giant Impact
Sean N. Raymond
Keywords
Planets
Definition
Synonyms
During planetary formation, a giant impact is a
collision between two planetary embryos with Jovian planets
Giant Planets 967
Definition Venus, the surface relief on the Moon, and the

multitude of stars which populate the Milky Way.
The four giant planets in our solar The discovery of the Galilean satellites had
system – ▶ Jupiter, ▶ Saturn, ▶ Uranus, and immense implications for the history of astron-
▶ Neptune – are found in the outer solar system, omy. It supported the heliocentric system pro-
at heliocentric distances ranging from 5 to 30 A- posed in 1543 by Nicolaus Copernicus and
U. Unlike the terrestrial planets – Mercury, promoted by Johannes Kepler, but still strongly
Venus, the Earth, and Mars – located in the attacked by the theologians of Rome. Later, Isaac
inner solar system, within 2 AU from the Sun, Newton elaborated the universal gravitation law
the giant planets are characterized by a large size, that provided the theoretical support of the helio-
a large mass, and a low density. They all have a centric model and opened the era of celestial
ring system and a large number of satellites. mechanics.
Since models show that the two largest giant Galileo also noticed the changing aspect of
planets have extensive gaseous envelopes, they Saturn as the orientation of the rings varied with G
are also referred to as “gas giants,” and there respect to the Earth, but he could not provide an
appear to be analogous objects in many extrasolar explanation. The answer was given by Christiaan
planetary systems. Tables 1 and 2 summarize the Huygens in 1659, who also discovered ▶ Titan,
main orbital and physical properties of the giant Saturn’s biggest satellite. At the end of the sev-
planets. enteenth century, large observatories were built
and planetary observations took place on a regu-
lar basis. ▶ Cassini observed the zone and belt
History structure of Jupiter and noticed its Great Red
Spot. He also discovered a gap inside Saturn’s
Early Exploration rings, now called the Cassini Division, and he
As naked-eye objects, Jupiter and Saturn have identified several icy satellites around Saturn.
been known since antiquity. Their astronomical Cassini also used the precise timing of the Gali-
observation started in 1610 when ▶ Galileo lean satellites’ mutual occultations to define a
Galilei used for the first time his new refractor universal time and allow navigators to determine
to look at celestial bodies. Galileo discovered the the longitude at sea. An even more spectacular
four big satellites which orbit around Jupiter result was the first measurement of the speed of
(later called Galilean satellites), the phases of light by Ole Rømer at Paris Observatory, based
Giant Planets, Table 1 Orbital properties of giant planets

Semimajor axis Inclination over the ecliptic plane Revolution period
Name (AU) Eccentricity ( ) (years)
Jupiter 5.20 0.054 1.30 11.86
Saturn 9.54 0.047 2.48 29.42
Uranus 19.2 0.086 0.77 83.75
Neptune 30.1 0.008 1.77 163.72
Giant Planets, Table 2 Physical properties of giant planets (mass and radius relative to Earth)
Name Mass (M
) Equatorial radius (R
) Density (g/cm3) Rotation period (h) Obliquity ( )
Jupiter 317.9 11.21 1.33 9.925 3.08
Saturn 95.16 9.45 0.69 10.656 26.73
Uranus 14.53 4.00 1.32 17.24 97.92
Neptune 17.14 3.88 1.64 16.11 28.80
968 Giant Planets
on the precise timing of the Galilean satellite from ground-based observations, thanks to the
occultations as a function of the Jupiter-Earth development of infrared and millimeter
distance. spectroscopy.
Thanks to improved techniques in the building
of glasses and lenses, astronomical telescopes got The Space Exploration of the Giant Planets
more and more powerful. William Herschel, in The space exploration of the giant planets started
particular, specialized in manufacturing a new in the 1970s with two programs, Pioneer and
generation of telescopes. In 1781, he discovered Voyager. Two spacecraft were first sent by
a new planet, which he called Uranus. The new NASA toward Jupiter and Saturn, Pioneer
planet was twice as far from the Sun as Saturn. At 10 and 11. Pioneer 10 was launched in 1972 and
the same time, as a natural consequence of New- flew by Jupiter at the end of 1973; Pioneer 11 was
ton’s theory of universal gravitation, a new sci- launched in 1973, flew by Jupiter in 1974, and
ence, celestial mechanics, was developing took advantage of Jupiter’s gravitational assis-
rapidly. In 1821, the first precise calculations of tance to encounter Saturn in 1979. The Pioneer
the giant planets’ orbits showed that the orbit of spacecraft were equipped, in particular, with a
the new planet was perturbed by another more camera, ultraviolet and infrared photometers, a
distant object. Two astronomers, John Couch magnetometer, a plasma analyzer, and particle
Adams in England and Urbain Le Verrier in detectors. The Pioneer mission provided us with
France, independently determined the position the first spectacular images of Jupiter’s atmo-
of the new object. John Adams was the first one spheric structures and Saturn’s rings, and they
to find the solution, but could not convince his explored for the first time the planets’
director about the importance of his discovery. In ▶ magnetospheres.
1846, Le Verrier sent the position of the new The first discoveries by the Voyager mission
planet to a German astronomer, Johannes Galle, appeared soon after the success of the Pioneer
who immediately found it within 1 of its spacecraft. Two identical spacecraft, Voyager
predicted position. The planet was called Nep- 1 and 2, were launched by NASA in 1977. Flybys
tune; this discovery marked the triumph of the of Jupiter took place in 1979, and those of Saturn
celestial mechanics. occurred in 1980 and 1981, respectively. Voy-
Visual observations and drawings remained ager 1 approached Titan for a close encounter of
for long the only means for planetary cartogra- Saturn’s satellite, while Voyager 2 used Saturn’s
phy. Photographic plates were used from the gravitational assistance to encounter the other
beginning of the twentieth century until the appa- giant planets. Voyager 2 flew by Uranus in 1986
rition of the ▶ CCD digital cameras in the 1980s. and by Neptune in 1989. The Voyager mission
In parallel, spectroscopic observations started to has been a spectacular success. In particular,
develop during the twentieth century. Spectro- Voyager 1 discovered the complexity of Jupiter’s
scopic observations are precious as they give dynamical atmospheric structure, the active vol-
information on the nature of the atmospheric canism of the inner Galilean satellite Io, the pos-
constituents of the planets. In 1932, methane sible water ocean below the surface of ▶ Europa,
and ammonia were detected in Jupiter’s the multiple structures of Saturn’s rings, the mag-
atmosphere; in spite of their low relative netosphere of Saturn, and the nature of ▶ Titan’s
abundance with respect to hydrogen, they were atmosphere. Voyager 2 discovered the magneto-
detectable because they are very active spectro- spheres of Uranus and Neptune, the unexpected
scopic agents. Methane was detected also on the tectonic activity of Uranus’ satellite ▶ Miranda,
other giant planets. Molecular hydrogen, the fascinating blue color of Neptune’s atmo-
although by far the dominant atmospheric sphere, and the active ▶ cryovolcanism at ▶ Tri-
species on all giant planets, was not ton’s surface. The Voyager mission marked a
detected before the 1960s. Since the 1970s, sev- major step in our understanding of the giant
eral other minor species have been detected planets (Fig. 1).
Giant Planets 969
Giant Planets, Fig. 1 The

Great Red Spot of Jupiter as
revealed by the Voyager
1 spacecraft in 1979
(# NASA)
After the flybys, a more in-depth exploration the atmosphere of Saturn. The exploration goes
of the giant planets required the launch of addi- on and will continue until 2017, with the
tional orbiters and probes. In 1989, the ▶ Galileo extended Cassini mission. The next step in the
mission was launched by NASA; it included an space exploration of the giant planets is the Juno
orbiter for successive flybys of the planet and the mission, launched by NASA in 2012, which will
Galilean satellites and a probe for in situ mea- encounter Jupiter in 2016 and explore its interior
surements of the Jovian atmosphere. In spite of a for a better understanding of its formation pro-
technical problem which disabled the large cesses. Later, the JUICE (Jupiter and Icy moons
antenna and imposed a very low telemetry rate, Explorer) mission, selected by ESA for a launch
the Galileo mission was also a great success. On in 2022, will approach the Jovian system in 2030.
December 7, 1995, the Galileo probe entered the After a series of flybys, the spacecraft will be put
Jovian atmosphere and transmitted data down to a in orbit around Ganymede for an exploration of
pressure level of 22 bars. The Galileo orbiter possible habitability conditions in the icy satel-
monitored the planet and the Galilean satellites lites (Fig. 2).
until 2003. Unexpectedly, Galileo’s magnetome- In addition to in situ exploration by dedicated
ter discovered, in particular, an intrinsic magnetic planetary spacecraft, one should not forget the
field on Ganymede. role of Earth-orbiting observatories. Both the
After Jupiter, Saturn and its system also International Ultraviolet Explorer (IUE),
deserved an in-depth exploration. In 1997, the launched in 1978, and the Hubble Space Tele-
ambitious ▶ Cassini mission, jointly developed scope (HST), launched in 1989, obtained UV
by NASA and ESA, was launched. The orbiter, spectra of the giant planets and Titan. The HST,
led by NASA, was designed for an in-depth mon- in addition, gave excellent quality images of the
itoring of Saturn, its rings, and satellites through giant planets and the outer satellites, allowing
multiple flybys. It approached Saturn’s system in long-term monitoring of the disk morphologies.
2004. On January 14, 2005, the ▶ Huygens The ▶ Infrared Space Observatory, launched by
probe, led by ESA, successfully landed on Titan’s ESA in 1995, sent infrared spectra which led, in
surface and sent to the world the first images of particular, to the detection of an external oxygen
this new world. Among other spectacular results, source on all giant planets and Titan. ISO was
the Cassini orbiter discovered lakes of hydrocar- followed by Spitzer, launched by NASA in 2003,
bons at the surface of Titan, cryovolcanism on and more recently by Herschel, launched by ESA
Enceladus, and a very complex meteorology in in 2009, which has pursued the far-infrared and
970 Giant Planets
the Sun’s rotation. The baselines of this model are

still valid today; later observations are totally
consistent with this scenario. First, the chemical
analysis of extraterrestrial samples (lunar sam-
ples and meteorites) has shown that all solar
system bodies formed some 4.56 billion years
ago, contemporaneously with the Sun itself. Sec-
ond, observations of young nearby stars have
revealed that the formation of a protoplanetary
disk, following the collapse of an interstellar
rotating cloud, is a common scenario of stellar
formation. In addition, hundreds of extrasolar
planets have been discovered around nearby
stars, either in these protoplanetary disks or
after the dissipation of these disks.
The main lines of the solar system formation
scenario can thus be summarized as follows. The
Giant Planets, Fig. 2 Saturn’s rings, as seen by the
Cassini spacecraft (# NASA)
protosolar disk, after the collapse of the
protosolar cloud, is mostly composed of gas
(hydrogen and, to a lesser extent, helium) with a
submillimeter exploration of the outer solar small fraction of dust (metals, silicates). At the
system. center of the disk, matter accretes to form the
young Sun. The temperature is high at the center
and decreases outward. The protosolar disk (see
Overview “▶ Solar Nebula”) is very turbulent such that
solid particles are permanently subject to mutual
The Formation of the Giant Planets collisions. Because all particles revolve around
The striking differences between the main proper- the center, their relative velocities are small and
ties of the giant planets, as described above collisions are not always disruptive; some of
(Tables 1 and 2), and the terrestrial ones suggest them lead to the formation of small aggregates.
a different formation scenario for the two types of Following a mechanism that is not presently fully
planets. Indeed, there is presently a general agree- understood (possibly helped by the turbulence),
ment within the scientific community about a for- some of these aggregates reach the kilometer
mation model of the solar system that does account size; they are called planetesimals. Then the big-
for these specific properties. gest fragments are able to capture the surrounding
It is now generally accepted that the solar particles by collisions and then by gravity.
system formed from a rotating fragment of an Numerical dynamical simulations show that
interstellar cloud, which collapsed into a disk after a few million years, a small number of big
perpendicular to its rotation axis. This model, objects (of the size of terrestrial planets and sat-
called “model of the primordial nebula,” was ellites) are formed.
first proposed in the eighteenth century by Why do we find two classes of planets, the
Immanuel Kant and later by Pierre-Simon de terrestrial and the giant ones? This separation is
Laplace. Their justification for such a model a result of the condensation sequence that takes
was the simple observation of planetary orbits: place within the disk as a function of the helio-
they all rotate around the Sun on nearly coplanar centric distance. Within the disk, the element
and concentric orbits, in the same direction abundances follow the universal cosmic rule:
(direct, i.e., counterclockwise as seen from the the lightest one (hydrogen) is the most abundant
north ecliptic pole), which is also the direction of (75 % per mass), followed by helium (23 %), and
Giant Planets 971
the 2 % left is made of all heavier elements. disk was warmer, the snow line was probably
Within these 2 %, the lightest (C, N, O) are located at about 3–4 AU from the Sun.
relatively more abundant than the heavier ones The “core accretion model” of the giant
(Si, Mg, metals). planet’s formation described above gives a natu-
As planets were formed from solid material, ral explanation for the basic properties of the
the nature of the planet depends upon the nature giant planets (size, mass, density, number of sat-
of the solid material available to form the plane- ellites). As will be discussed below, another
tesimals. In the protosolar disk, the temperature observational fact supports this model: the ele-
decreases as the heliocentric distance increases. mental and isotopic abundances measured in the
Two cases can be identified: giant planets.
Close to the Sun, within a few AU, the tempera- Gaseous Giants and Icy Giants
ture (above 200 K) was such that the only solid Table 2 shows the giant planets fall into two
matter available was made of heavy atoms distinct subclasses. With masses of about G
(metals, silicates). Because heavy elements 300 and 100 terrestrial masses, Jupiter and Saturn
are not abundant in the disk, the accretion are mostly composed of protosolar gas: they are
process led to the formation of relatively called the gas giants. In contrast, Uranus and
small and dense planets: the terrestrial planets, Neptune, with masses around 15 terrestrial
also called the rocky planets. masses, are mostly composed of metal, rock,
At larger heliocentric distance, the temperature and water, methane and ammonia compounds,
(below 180 K) became low enough for the the latter mostly in gaseous states. They are called
small molecules (H2O, CH4, NH3, H2S, CO2, the icy giants.
etc.) to condense. As these species were much What can be the reason for such a difference?
more abundant than the heavier elements, they No definite answer can be given presently, but a
together with metals and silicates accreted into plausible explanation can be proposed. Jupiter,
bigger cores, able to reach 10–15 Earth formed just beyond the snow line, benefited from
masses. At this point, models predict that a maximum of icy material and a maximum of
their gravity field is sufficient for the sur- surrounding gas. To a lesser extent, Saturn’s for-
rounding nebula to collapse in a disk, in the mation followed the same way. Jupiter and Sat-
equatorial plane of the planetary core. As the urn probably completed their formation within a
nebula is mostly made of hydrogen and few million years at most. In contrast, Uranus and
helium, the planets formed after these col- Neptune, formed at larger heliocentric distances,
lapses are very big, with a low density: they needed a longer time to accrete their icy core.
are the giant planets with their ring and Possibly, they reached the critical mass of 10 ter-
satellite systems, formed within their restrial masses late enough for the protosolar disk
equatorial disk. to have dissipated, leaving little gaseous material
for the planets to continue accretion. The lifetime
Between the terrestrial and giant planets, the of the protosolar disk, as inferred from the obser-
line of ice condensation is called the snow line. vation of protoplanetary disks around nearby
Water plays a special role for two reasons: first, stars, was probably not longer than 10 million
formed from two abundant atoms, H and O, water years, which may have been the time required
is, among the ices, the most abundant small mol- by the icy giants to accrete their icy core.
ecule; second, it is the first molecule to condense
as the temperature decreases. Other ices (NH3, Migration of the Giant Planets
CO2, CH4, etc.) condense at greater heliocentric Recent discoveries of extrasolar planets suggest
distances. Presently, water condenses at about that migration has been a common phenomenon
2 AU from the Sun, as shown by cometary activ- in the planetary disks of nearby stars. Migration is
ity. At the time of planets’ formation, when the usually generated by the interaction of the planet
972 Giant Planets
with the turbulent protoplanetary disk and leads the expected to be found in reduced form. This is the
planet to move inward, from the outer stellar sys- case for methane and hydrocarbons produced by
tem to the close environment of its star. In the case its photodissociation. In addition, NH3, PH3,
of the solar system, dynamical simulations also H2O, GeH4, and AsH3 are observed in Jupiter’s
suggest some migration might have taken place. and Saturn’s tropospheres; these species are not
According to the “Nice Model” developed by observable on Uranus and Neptune because they
Alessandro Morbidelli and his colleagues, after condense below the observable atmospheric
the dissipation of the protosolar disk, the giant levels (at pressure levels of tens of bars). Methane
planets were located on quasi-circular orbits also condenses on Uranus and Neptune at a level
between 5.5 and 17 AU, i.e., significantly closer of 1 bar (T = 80 K) but, especially on Neptune,
to the Sun than now. New numerical simulations its stratospheric content is sufficient to lead to the
suggest that this special configuration was the formation of hydrocarbons (in particular C2H2
result of an earlier migration (the so-called and C2H6). The atmospheric composition of the
Grand Tack) during which Jupiter came inward giant planets is listed in Table 3.
as close as Mars’ orbit (thus interrupting the Other unexpected stratospheric species have
growth of Mars, which would explain the low been discovered. In 1992, ground-based millime-
mass of the planet) and then migrated outward ter observations of Neptune led to the detection of
under the influence of Saturn. Later, gravitational ▶ stratospheric CO and HCN, in abundances
perturbations may have induced again a moderate much larger than the predicted ones (those
motion of Jupiter toward the inner solar system, predicted values were actually observed in Jupi-
while the three other giant planets moved out- ter’s and Saturn’s stratospheres). In 1997, the ISO
ward. During this motion, the Jupiter-Saturn sys- satellite discovered the presence of H2O and CO2
tem crossed the 2:1 resonance (Saturn’s on all giant planets and Titan (only CO2 on Ura-
revolution period being twice Jupiter’s). nus). CO was detected on Uranus from ground-
According to dynamical simulation, this event based infrared spectroscopy. H2O and CO were
generated a great perturbation in the inclinations later also studied by the Herschel satellite.
and ellipticities of solar system small bodies. Because the thermal vertical profiles of the giant
A clear signature would be the “▶ Late Heavy planets exhibit a cold trap at the level of the
Bombardment” (LHB), traces of which are minimum temperature, at a pressure level of
observed on the surfaces of all bare solar system 0.1 bar (see below), the stratospheric oxygen
objects, from Mercury and the Moon to the outer species had to be of external origin. The source
satellites. Crater rate counting on these objects might be either local (rings and satellites) or
shows that the event took place some 3.8 billion interplanetary (comets or flux of micrometeor-
years ago, i.e., about 800 million years after the ites). Following Herschel observations, the
planet’s formation. water source is believed to be the Shoemaker-
Comparison with other planetary systems Levy collision of 1994 for Jupiter and the
raises a question: why was migration moderate ▶ Enceladus torus in the case of Saturn.
in the solar system, while it seems to have been From the abundances of ▶ tropospheric spe-
very efficient around nearby stars? Possibly, this cies, elemental and isotopic ratios have been
was the result of the specific masses and locations determined. These parameters are important as
of Jupiter and Saturn at the beginning of the they provide precious constraints for the forma-
scenario, but we have presently no firm answer tion models of the giant planets. Indeed,
to this question. according to the nucleation formation model
described above, heavy elements are expected
Atmospheric Composition of the Giant to be enriched with respect to protosolar values;
Planets the larger the mass fraction of the icy core, the
Because the atmospheres of the giant planets are larger the enrichment. A simple model based on
dominated by hydrogen, atmospheric species are an initial icy core of 12 terrestrial masses predicts
Giant Planets 973
Giant Planets, Table 3 Giant planets: mixing ratios of atmospheric constituents (fractional number density per unit
volume relative to H2)
Constituent Jupiter Saturn Uranus Neptune
Troposphere
H2 1 1 1 1
He 0.157 0.13 0.18 0.23
CH4 2.1103 4.4103 2102 4102
NH3 2104 2–4104
PH3 6107 1.7106
H2O 1.4105 2107
GeH4 71010 2109
AsH3 31010 2109
CO 1.5109 2109 3108 106
Stratosphere
CH4 2.1103 4.4103 3105–104 7104
G
C2H2 3108 2107 2107 3107
C2H4 7109 a
107
C2H6 2106 3106 2106
CH3 5108 5108
C3H4 a
61010
C3H8 6107
C4H2 a
91011
9 a
C6H6 210
H2O 3109 108 7109 2109
CO 1.5109 2109 3108 106
CO2 31010 31010 51010
HCN 31010
H3+ a a a
13 15 12 13
NB: Isotopic species have been also detected: HD, CH3D, CH4, NH3, C CH2
a
Detected
enrichment in heavy elements by a factor of 4 for problem. It is hoped that the Juno mission will
Jupiter, 9 for Saturn, and about 30 for Uranus and bring new answers to this question by measuring
Neptune, which is in full agreement with the the O/H ratio in the planet’s interior.
carbon enrichment measured from tropospheric Another additional measurement supports the
methane in the four giant planets. It is also fully nucleation model. The D/H ratio measured in the
consistent with the enrichments measured in situ outer solar system is diagnostic of its formation
by the mass spectrometer of the Galileo probe for temperature. Indeed, ▶ deuterium is enriched
several elements (C, N, S, Ar, Kr, Xe). These with respect to hydrogen in ices, due to
results provide a clear validation of the nucle- ion-molecule and molecule-molecule reactions,
ation formation model. Still, the Galileo mea- as observed in the laboratory and in the ▶ inter-
surements on Jupiter raise a problem: all stellar medium. The observed D/H ratio increases
elements seem to have been equally trapped in as the temperature decreases. The D/H ratio has
ices; but N and Ar cannot be trapped unless at been measured in the giant planets by the ISO
very low temperature (<40 K). This suggests that satellite from HD infrared lines. These lines were
the planetesimals which formed Jupiter were later reanalyzed on Uranus and Neptune by the
accreted at very low temperatures, much lower Herschel satellite. In the case of Jupiter and Sat-
than expected at Jupiter’s orbit. The origin of urn, D/H has been found to be in agreement with
Jupiter’s planetesimals is presently an open the protosolar D/H value, which is consistent
974 Giant Planets
with the fact that they are mostly made of pressure levels of 5 bar (Jupiter) and 10 bars
protosolar gas. In contrast, following Herschel’s (Saturn). In the case of Uranus and Neptune,
results, D/H in Uranus and Neptune is enriched according to the models, CH4 is expected to con-
by a factor of about 2–2.5, which corroborates dense at a level of 1 bar (T = 80 K). Hydrocar-
that these planets have an icy core. bon hazes (especially C2H6) should be present in
The abundance of helium in the giant planets the stratosphere. A H2S cloud could be present at
is also diagnostic of their formation and evolution about 3 bars (T = 120 K). In the lower tropo-
processes. The He mass fraction Y was measured sphere, NH4SH and H2O clouds could be present
by the Voyager IRIS infrared spectrometers and, at temperatures of 240 K and 270 K, respectively,
in the case of Jupiter, by the Galileo probe. at pressure levels of several tens of bars.
Results were Y = 0.234 +/0.05 for Jupiter, How do the observations fit the models? The
0.21+/0.3 for Saturn, 0.26 +/0.05 for Uranus, spectroscopic identification of condensates is
and 0.32 +/0.05 for Neptune. For Uranus and much more ambiguous and difficult than the
Neptune, the helium abundance is consistent with detection of gaseous species. Still, imaging spec-
the protosolar abundance (Y = 0.275 +/0.01). troscopy at different wavelengths provides a 3D
In the case of Saturn and Jupiter, the apparent imaging of planetary disks. Typically, ultraviolet
depletion has been interpreted as a possible con- radiation probes the stratosphere, while the visi-
densation of helium within the liquid hydrogen ble spectrum probes the upper troposphere; the 5-
ocean in the planets’ interiors (see below). micron region, known as an “atmospheric win-
dow,” probes the troposphere down to a pressure
Thermal Structure, Cloud Structure, of a few bars.
and Dynamics Images of Jupiter and Saturn in the visible
The atmospheres of all giant planets are charac- range show the typical zone-belt structure char-
terized by several layers: (1) a convective ▶ tro- acteristic of a Hadley-type convective circulation
posphere where the temperature decreases as the in a fast rotating object. Zones are cloudy regions
altitude increases following an adiabatic lapse of ascending motion, while belts are dry, cloud-
rate. The minimum temperature is reached at free regions of subsidence. The yellow color of
the tropopause, at a pressure level of about Jupiter’s zones is consistent with the presence of
100 mbar for all giant planets. The temperature an ammonia cloud. In Saturn, the more uniform
at this level is 110 K for Jupiter, 90 K for Saturn, yellow color of the disk implies a thicker
and 50 K for Uranus and Neptune. Above the ammonia cloud, consistent with the colder envi-
tropopause, in the ▶ stratosphere, the tempera- ronment of the planet. On Jupiter, dark-blue
ture increases again with altitude. In the lower regions in the belts have been attributed to water
stratosphere, the heating mechanism is probably clouds. The Great Red Spot, known for over three
the absorption of the solar flux by methane and centuries, is believed to be a giant anticyclonic
hydrocarbon aerosols formed by methane photo- structure. Its stability over such a long time is still
dissociation. At higher altitudes, other heating a puzzle. Its red color has not been unambigu-
processes must be involved, such as gravity ously interpreted. It could be due to the presence
waves and energy deposition by high-energy of sulfur or phosphorus compounds or to hydro-
particles. carbon polymers formed from the irradiation of
The cloud structure of the giant planets can be ices by high-energy particles. In Saturn, a huge
estimated on the basis of thermochemical equi- storm developed in the northern hemisphere in
librium models. In the case of Jupiter and Saturn, December 2010. This phenomenon has been
a cloud of ammonia is expected at a level of observed for over a century with a period of
0.5–1 bar, respectively (T = 145 K). A NH4SH approximately 28 years near the northern summer
cloud is expected at T = 210 K, at pressure levels solstice. It is attributed to a massive atmospheric
of 2 bars (Jupiter) and 4 bars (Saturn). At deeper upwelling generated by a seasonal insolation
levels, a H2O cloud should occur at 270 K, at increase.
Giant Planets 975
The Galileo and Cassini observations have had disappeared a few years later when Neptune
shown that, on both Jupiter and Saturn, the mete- was imaged by the HST. Many other variable
orology is by far more complex than described by bright and dark spots, at different altitudes, have
a simple Hadley-type convective model. In the been monitored over the years. White spots are
case of Jupiter, the Galileo probe entered a region believed to be high-altitude cirrus of methane.
especially dry and free of clouds. In particular,
the cloud structure described above was not Internal Structure
found, and the measured water abundance was Our knowledge of the giant planets’ ▶ interior
much lower than expected. This measurement structure relies on theoretical models, constrained
was not considered as representative of the O/H by a few physical parameters (mass, diameter,
value in the deep troposphere of the planet. In the density, gravitational moments). According to
case of Saturn, near-infrared images revealed a these models, the internal structures of Jupiter
huge cyclone at the South Pole and a complex and Saturn consist of three regions: a central core
hexagonal structure surrounding the North Pole. of ices and rocks, an ocean of metallic hydrogen, G
In contrast with the other giant planets, the and an envelope of molecular hydrogen. At the
image of Uranus shows little contrast. The center, the temperature is expected to be
images taken by Voyager 2 in 1986 showed no 23,000 K (Jupiter) and 12,000 K (Saturn); the
longitudinal structure. At the time of the Voyager pressure is estimated to 50–100 Mbar (Jupiter)
encounter, the rotation axis of Uranus was nearly and about 50 Mbar (Saturn). In the case of Uranus
aligned with the Sun-planet axis. Twenty years and Neptune, the models predict also a core made
later, when Uranus came to equinox, HST images of ices and rocks but no metallic hydrogen. The
revealed more activity, with cloud features at low core would be surrounded by a mixture of ices and
latitude. In spite of some possible seasonal activ- then by an envelope of molecular hydrogen. The
ity, Uranus is much less dynamically active than central temperature would be about 7,000 K and
the three other giants. Is this difference connected the central pressure about 6 Mbar.
to the special orientation of its rotation axis, very Other information relative to planetary inte-
close to the ecliptic plane, or is it connected with riors come from the internal energy source that
the absence of an internal energy source (see has been measured by Voyager from the total
below)? Or are the two effects connected with infrared emission of the planets. An internal
each other? This is an open question. energy source has been detected on Jupiter, Sat-
Finally, Neptune exhibits a strong dynamical urn, and Neptune; its ratio to the absorbed solar
activity, which is especially remarkable in view energy amounts to 1.67 for Jupiter, 1.78 for Sat-
of its large distance from the Sun. Part of the urn, and 2.6 for Neptune. In the case of Uranus,
explanation may come from its strong internal the ratio was less than 1.1; no internal energy was
source of energy (see below). Methane in Nep- detected. A plausible source for the internal
tune is supersaturated, which induces a strong energy is radiative adiabatic cooling, following
methane photochemistry. The explanation might the early accretion phase of the planet’s forma-
be the continuous insolation of the South Pole tion. If it is indeed the case, then the absence of
over the past 40 years, which could be responsi- internal energy on Uranus is puzzling. In the case
ble for the temperature increase measured at the of Jupiter and Saturn, according to models, an
tropopause; methane would be released in the additional internal source might come from the
stratosphere from the South Pole and spread condensation of helium inside the metallic ocean
over the planet. An active meteorology has been of hydrogen. Helium droplets would sink toward
observed on Neptune over the past decades. In the center, releasing energy upward and contrib-
1989, at the time of the Voyager encounter, a uting to the depleted helium abundance in the
large feature (the Great Dark Spot) was observed. outer layers; no helium condensation would be
It was interpreted as a giant anticyclone, similar expected in the case of Uranus and Neptune, as
to Jupiter’s Great Red Spot. However, the feature the internal pressure is not sufficient for hydrogen
976 Giant Planets
to be in liquid form. This process would be con- rotation axis by a large angle (59 and 47 , respec-
sistent with the helium measurements in the giant tively), with an offset of 0.3 and 0.5 planetary radii
planets (see above). with respect to the planet’s center. This configura-
tion induces complex magnetospheric structures,
Magnetospheres modulated by the rotation period of the planets. In
Like the Earth, all giant planets have a ▶ magne- the case of Uranus, the situation is even more
tosphere. In the case of Jupiter, its magnetic activ- complicated due to the position of the rotation
ity was first identified in 1955 by the detection of axis, close to the ecliptic plane, which leads to
strong radio emission. In addition to the thermal strong seasonal effects.
emission of the planet, nonthermal emission was
found at centimeter and decimeter wavelengths. It Rings and Satellites
was attributed to synchrotron emission of electrons All giant planets have a ring system (see “▶ Plan-
trapped in the radiation belts of a newly detected etary Rings”) and a large number of satellites; as
magnetic field. The magnetospheres of Jupiter and mentioned above, this is a direct consequence of
Saturn were first explored by Pioneer 10 and 11 in their formation scenario. The satellites were
1974 and 1979, respectively, and the magneto- formed within the planet’s equatorial plane,
spheres of Uranus and Neptune were discovered after the collapse of the surrounding nebula on
by Voyager 2 in 1986 and 1989, respectively. the initial icy core.
The magnetic fields of the giant planets, to first Rings are mostly found within the ▶ Roche
order, can be approximated by a dipole, probably limit, at about two planetary radii from the
generated by motions within the liquid central planet’s center; in this region, tidal forces prevent
nucleus, also activated by the fast rotation of the the accretion of satellites, and dust particle
planets. revolve along circular orbits around the planet.
The magnetospheres of the giant planets show The giant planets show very different ring
a structure comparable with the terrestrial one systems. Saturn’s rings have been observed over
and result from the interaction of the magnetic four centuries, while the three other ring systems
field with the ionized particles of the solar wind. were still undetected a few decades ago. The ring
In the case of Jupiter, the inner magnetosphere, system of Jupiter, made of three tenuous compo-
within 20 planetary radii, has a dipole structure, nents, was discovered by Voyager 1. They are
inclined by 11 with respect to the rotation axis, made of micron-sized particles, probably ejected
with radiation belts similar to the Van Allen belts from the small nearby satellites (Thebe,
on Earth, where energetic particles are trapped. Amalthea, Metis, and Adrastea). These particles
The magnetic field is perturbed by Jupiter’s sat- have a very short lifetime, due to dynamical
ellites Io, Europa, and Ganymede. Aurorae are effects and erosion.
observed in the ionosphere at high latitude, in the The ring systems of Uranus and Neptune were
form of localized emissions in the ultraviolet and both discovered by ground-based stellar ▶ occul-
in the near-infrared range. tation measurements. In 1977, nine very narrow
The Jovian magnetic field (14 G at the North rings were discovered around Uranus. The obser-
Pole, 4 G at the equator) is ten times stronger than vation was made easier by the fact that the rota-
the terrestrial magnetic field. The other giant tion axis of Uranus was almost facing the Earth at
planets have significantly weaker magnetic fields that time. The rings of Uranus were later imaged
(0.2–0.3 G). Saturn’s magnetosphere shows no by Voyager 2 and then by the ▶ HST. In 1984,
radiation belts, because of the presence of the another stellar occultation experiment was
rings. As Saturn’s magnetic axis is almost aligned performed on Neptune. Absorption of the stellar
with its rotation axis, its magnetosphere is much flux was detected on one side of the planet only; it
more regular than the Jovian one. Both Uranus and was interpreted as the presence of incomplete
Neptune show a very strange configuration, as rings or “arcs.” In 1989, Voyager 2 confirmed
their magnetic axes are tilted with respect to their this result but found that the rings were actually
Giant Planets 977
Giant Planets, Fig. 3 The rings of Neptune as observed

by Voyager 2 in 1989. The denser regions correspond to
the “arcs” previously identified from ground-based solar
occultation observations in 1984 (# NASA)
complete, but denser in some locations. The rings Giant Planets, Fig. 4 The system of Uranus, observed in
the near-IR range at the VLT (European Southern Obser-
of Uranus and Neptune have a low albedo, pos- vatory: ESO) in 2002. On this picture, satellites and rings
sibly due to the presence of organic polymers are overexposed and the planet is dark because the near-IR
resulting from the irradiation of organic ices by filter (K-band) corresponds to a strong absorption band of
high-energy magnetospheric particles (Figs. 3 methane. North is at the bottom right and east is at the top
right. The satellites, from left to right, are Titania,
and 4). Umbriel, Miranda, Ariel, and Oberon (# ESO)
The detailed structure of Saturn’s ring system
has been revealed by Pioneer and Voyager and is
being extensively explored by the ▶ Cassini satellites are made of ices. Water dominates in
orbiter. Apart from the well-known A, B, and the case of Jupiter (with the exception of Io,
C main rings, a thin ring (D) has been found which has lost its water due to its active volca-
closer to the planet; other rings (A, F, G, E) nism) and Saturn. The satellites of Uranus and
have been found outside. The Cassini observa- Neptune, which have a lower albedo, probably
tions have shown that the E-ring is fed with the contain other types of ices (N2, CH4, CO2, etc.) as
matter ejected by ▶ Enceladus, which shows evi- is observed on ▶ Triton.
dence for active ▶ cryovolcanism. The radial distribution of satellites shows
large variations from one planet to another.
Outer Satellites There are some analogies between the Galilean
All giant planets have many satellites, either reg- satellites of Jupiter, which extend from 6 to
ular (i.e., formed in the equatorial plane of the 26 planetary radii, and the five big satellites of
planet) or irregular (i.e., resulting from the cap- Uranus, which extend from 5 to 23 planetary
ture of a distant asteroid or a ▶ trans-Neptunian radii. In contrast, Saturn’s largest satellite,
object). All regular satellites are in synchronous ▶ Titan, is located at 20 planetary radii, beyond
rotation with their planet. five midsized satellites. Titan is unique in having
The outer satellites are all different, and each a dense neutral atmosphere. Neptune’s system is
of them deserves a specific description. Here we different, because of the capture of the big satel-
will just mention some general trends. All outer lite ▶ Triton, located at 14 planetary radii.
978 Giant Planets, Model Atmospheres of
No midsized satellite is found at Neptune except

the small Proteus at five planetary radii and the Giant Planets, Model
very distant Nereid. Triton’s capture might have Atmospheres of
expelled other regular satellites previously
formed in Neptune’s equatorial plane. ▶ Exoplanets, Modeling Giant Planets’
Atmospheres
See Also
▶ Galileo Galilei
▶ Huygens Giga-Annum
▶ Interior Structure, Planetary
▶ Jupiter ▶ Ga
▶ Neptune
▶ Planetary Rings
▶ Saturn Gigayear
▶ Uranus
▶ Ga
(1979) Mission to Jupiter and its satellites (Voyager

1 encounter). Science 204:945–1008
Giotto Spacecraft
(1996) Galileo at Jupiter: results from the Orbiter. Science
274:377–412 Anny-Chantal Levasseur-Regourd
(1996) Galileo at Jupiter: results from the Probe. Science UPMC University of Paris 6/LATMOS-IPSL,
272:837–860
Bagenal F, Dowling T, McKinnon W (2007) Jupiter: the
Paris, France
planet, satellites and magnetosphere. Cambridge Uni-
versity Press, Cambridge Planetary Science, New
York Keywords
Bergstrahl JT, Miner ED, Matthews MS (1990) Uranus.
University of Arizona Press, Tucson
Bézard B, Drossart P (2001) Neptune. In: Murdin Comet; Nucleus; Coma; Water; Dust; Organics;
P (ed) The encyclopedia of astronomy and astrophys- CHON; Giotto; Giotto extended mission; Space-
ics. IoP, Bristol, pp 798–1806 craft; ESA; Halley; Grigg-Skjellerup
Cruikshank DP (1995) Neptune and triton. University of
Arizona Press, Tucson
Drossart P (2001) Saturn. In: Murdin P (ed) The encyclo-
pedia of astronomy and astrophysics. IoP, Bristol Definition
Encrenaz T (2001) Jupiter. In: Murdin P (ed) The ency-
clopedia of astronomy and astrophysics. IoP, Bristol,
pp 1330–1336
The Giotto spacecraft (Fig. 1), the first ESA
Gehrels T, Matthews MS (1984) Saturn. University of (European Space Agency) interplanetary probe,
Arizona Press, Tucson was designed to flyby ▶ comet Halley. Launched
Herbert F (2001) Uranus and Neptune: atmospheres, ion- on 2 July 1985 by an Ariane-1 rocket from
ospheres ad magnetospheres. In: Murdin P (ed) The
encyclopedia of astronomy and astrophysics. IoP,
Kourou, Giotto succeeded in approaching the
Bristol, pp 3408–3415 cometary nucleus to within 600 km on
Lunine JI (1993) The atmospheres of Uranus and Neptune. 14 March 1986. Through its first accurate images
Annu Rev Astron Astrophys 31:217–263 of a nucleus and in situ studies of gases and dust
Miner ED (ed) (1998) Uranus – the planet, rings and
satellites. Wiley, New York
particles within a coma, the mission has revealed
Smith BA et al (1986) Voyager 2 in the uranian system: the complexity of comets. Afterwards, the Giotto
results from the imaging team. Science 233:97–102 spacecraft was re-oriented in order to study comet
Giotto Spacecraft 979
European mission, provided a close-up view of

the cometary nucleus, afterwards described as an
icy dirt ball, together with evidence for previ-
ously unsuspected properties of its dust particles,
found to be under-dense and rich in refractory
organics. Giotto was also the first mission to be
reactivated after hibernation and to return from
interplanetary space for an Earth swingby, which
oriented it toward a second comet.
Basic Methodology
At the end of the 1970s, a mission to a comet was G

mandatory to assess the existence of the nucleus,
the properties of the coma, and the solar wind
interactions. The expected return in 1985–1986
of the active short-period comet 1P/Halley, the
trajectory of which was well known, made it an
obvious target for a ballistic flyby. Its retrograde
orbit (with respect to that of the Earth and of a
space probe) nevertheless implied a huge relative
velocity between the comet and any spacecraft,
close to 70 km/s. In July 1980, once it had
become clear that a sophisticated ▶ NASA mis-
Giotto Spacecraft, Fig. 1 Giotto spacecraft sion with a flyby of Halley and a rendezvous with
another comet would not take place, the Euro-
pean Space Agency approved a mission specifi-
Grigg-Skjellerup, which was flown by on 10 July cally dedicated to the study of Halley during its
1992, at a nucleus distance in the 150–200 km passage in the ecliptic at the orbital node closest
range. to its perihelion, i.e., in March 1986.
The onboard experiments (Fig. 2) were
selected in January 1981. They consisted of ten
History hardware experiments with a total mass of about
60 kg, i.e., one camera, three mass spectrometers
Giotto was named after the painter Giotto di (neutral, ion, dust), one dust impact detector, one
Bondone, who, in 1301, depicted a comet as the optical probe, two plasma analyzers, one ener-
star of Bethlehem in his fresco Adoration of the getic particles analyzer, and one magnetometer,
Magi in Padua. The comet, easily visible from plus one radio-science experiment. The space-
Europe at that time, was later called 1P/Halley. craft had a cylindrical shape (diameter 1.86 m,
height 2.85 m, dry mass 574 kg), with its main
cylinder covered by a solar array. It was spin
Overview stabilized (period of 4 s), with the spin axis
aligned with the relative velocity vector during
Although comets had been observed for centu- Halley encounter. A two-stage aluminum-kevlar
ries, their knowledge remained limited until the dust shield, perpendicular to the spin axis,
1986 flybys of Halley by an armada of spacecraft protected Giotto against high-velocity dust
(see below). Giotto, the first independent impacts; the despun (to counteract the effect of
980 Giotto Spacecraft
MAG
F.M. Neubauer
Univ. zu koin, Germany
HMC
H.U. Keller
MPl, Lindau, Germany
DID MAG
J.A.M. McDonnell
Univ. of Kent.Canterbury, UK
RPA
H. Rème
CESR, Toulouse, France OPE
JPA
A. Johnstone
Mullard Space Sc Lab.
Holmbury St Mary. UK
PIA
J. Kissel
MPl. Heidelberg. Germany
OPE
A.C. Levasseur-Regourd
CNRS, Verrières-le-Buisson, France
DID HMC
J.A.M. McDonnell
Unv. of Kent.Canterbury. UK EPA
DID
EPA DID
S.M.P. McKenna-Lawlor
St. Patrick’s Coll. Maynooth, irl
NMS
D. Krankowsky RPA
MPI, Heidelberg, NMS
Germany
JPA
JPA
A. Johnstone
Mullard Space Sc Lab.
Holmbury St Mary. UK
IMS
H. Baisiger
JPA
Univ. of Bern, Switzerland
PIA
GRE
P. Edenhofer IMS
Ruhr University. Bochum, Germany
Giotto Spacecraft, Fig. 2 Giotto onboard experiments
the spin) parabolic reflector of the high-gain eccentric geosynchronous transfer orbit. About
antenna was mounted on a tripod at the other 1 day and a half later, its solid-propellant motor
end, in order to be oriented toward the Earth was fired at perigee to inject the spacecraft into a
during the flyby. solar ballistic orbit that allowed it, after a cruise
After 5 years of development and tests, Giotto of about 685 million kilometers, to flyby the
was launch by an Ariane-1 rocket from Kourou, cometary target without major trajectory adjust-
French Guyana, on July 2, 1985, into a highly ments. The encounter took place on March
Giotto Spacecraft 981
Giotto Spacecraft, Earth

Fig. 3 Geometry of Comet 44.3°
Halley flyby Bow s
hock
Contact
surface
107.2°
Sun
Spacecraft
trajectory
68.7 km/s
13–14, 1986, with a relative velocity of 68.4 km could be established that it had survived with
s1, at 0.89 AU heliocentric distance and minor degradation of the Halley flyby. On July
0.98 AU geocentric distance and for a phase 2, 1992, after six orbits around the Sun, Giotto
angle of 107.05 . The closest approach took was back in the vicinity of our planet at
place immediately after midnight UT, at 23,000 km altitude. The Giotto Extended Mission
(596 2) km distance from the nucleus (Fig. 3). (GEM) to a second cometary target had already
To allow Giotto to approach as close as possi- been approved by ESA, and the orbit of the
ble to the nucleus without being destroyed, a spacecraft was retargeted through an Earth grav-
unique international cooperation had been orga- ity assist toward 26P/Grigg-Skjellerup, a Jupiter
nized. An armada of spacecraft had been family comet.
launched toward Halley, including besides Giotto This last flyby took place on July 10, 1992,
the Japanese Suisei with a nucleus miss distance with a relative velocity of 14 km s1, at 1.01 AU
of 151,000 km and the Russian Vega 1 and Vega heliocentric distance and 1.43 AU geocentric dis-
2, respectively, at 8,890 and 8,030 km. The path- tance. While the Giotto scientific payload was
finder concept, developed in cooperation fully operational for the Halley flyby, it was
between ESA, IKI (USSR), and NASA (USA) only 50 % operational for Grigg-Skjellerup and
was used to define the photometric center no images were provided by the damaged HMC
(presumed to correspond to the nucleus) on the camera. Nevertheless, the OPE (Optical Probe
images collected by the Vegas, the positions of Experiment), undamaged at the rear of the space-
which were determined by the American Deep craft, derived a nucleus miss distance below
Space Network. The distance of closest approach 200 km, most likely in the 150–200 km range,
of Giotto was accurately controlled through this from the monitoring of the evolution of the
unique effort, allowing the spacecraft to approach brightness under the observational geometry.
the nucleus without being totally destroyed by Among the huge wealth of unique results,
impacts from dust particles hitting it with a high many are relevant from an astrobiology
relative velocity. perspective.
Giotto was actually hit by a rather large dust
particle 14 s before closest approach, leading by
nutation to a shift of the spacecraft angular Key Research Findings
momentum vector of 0.9 and to an intermittent
Earth data link for 32 min. Two weeks after the The Giotto HMC camera provided the first accu-
encounter, the spacecraft was put into hiberna- rate images of a cometary nucleus. They revealed
tion. After its reactivation in February 1990, it a slightly reddish irregular object, significantly
982 Giotto Spacecraft
water. Nevertheless, these comets are not origi-

nating in the Kuiper Belt, and water on Earth
might also come from icy small bodies formed
in the outer asteroid belt.
One of the most unexpected discoveries of
Halley’s missions was that the dust mass spec-
trometers (PIA on board Giotto, PUMA on Vega)
not only detected rock-forming elements (e.g.,
Mg, Si, Ca, Fe) but also revealed light elements,
so-called CHON material, most likely consisting
of complex polymeric organic molecules, com-
posed of carbon, hydrogen, oxygen, and nitrogen
(CHON), in agreement with data from one
plasma analyzer. Such refractory organics, the
presence of which was later confirmed for
comet Wild 2 by the ▶ Stardust mission, had
quite likely been formed in the interstellar
medium. Besides, the properties of dust particles
Giotto Spacecraft, Fig. 4 Image of Comet Halley were varying with distance to the nucleus and
nucleus obtained by Giotto with time after ejection, most likely under evap-
oration and fragmentation processes. Compari-
sons between the results of OPE and DID (Dust
darker and bigger (about 15 km long and 7.5 km Impact Detector) have shown that, on the aver-
by 8 km wide) than anticipated, with various age, the geometric ▶ albedo of the dust particles
topographic features (hills, ridges, craters, cliffs, was very low (about 0.04) and their density
ridge, central depression). Well-defined dust extremely reduced (about 100 kg m3),
structures with the appearance of narrow suggesting that they mostly consisted of fluffy
filaments were noticed in the inner coma, possi- aggregates.
bly originating from active regions covering 10 % During the Giotto Extended Mission to Grigg-
of the surface. The mass of the nucleus was Skjellerup, fragmentation processes could be
too low to be derived from the spacecraft suspected, with one event noticed by OPE at
trajectory perturbations near closest approach. about 1,000 km distance from the nucleus, tenta-
Nevertheless, estimations of the mass from tively interpreted by the presence of an active
non-gravitational effects (later validated fragment (10–100 m size) releasing dust in the
through the ▶ Deep Impact mission) and of the inner coma. All together, discoveries of
shape from imaging yield a density of (1) porous nuclei that may suffer fragmentation
550 250 kg.m3, typical of a highly porous processes and (2) refractory organic molecules in
object (Fig. 4). low-density dust particles indicate that comets
The icy component of Comet Halley mostly can significantly replenish the zodiacal cloud of
consisted of water. Independent analyses of the interplanetary dust with organics. During the
D/H ratio from NMS and IMS (Neutral and Ion ▶ Late Heavy Bombardment epoch, there was
Mass Spectrometers) lead to a value in cometary certainly a huge amount of interplanetary dust
water of (3.08 0.3) 104, i.e., twice that of particles of cometary origin. Their highly porous
seawater on Earth, corresponding to an enrich- structure, which leads to a significant decelera-
ment by a factor of 15 relative to the protosolar tion and heat transfer in planetary atmospheres,
cloud. With similar results later obtained for a could have contributed to extraterrestrial delivery
couple of other comets, it may indicate that of carbonaceous compounds within the atmo-
comets only marginally contributed to Earth’s spheres of terrestrial planets.
GJ 667C: First System with Multiple Super-Earths in the Habitable Zone 983
Future Directions Keller HU, Britt D, Buratti BJ, Thomas N (2004) In situ
observations of cometary nuclei. In: Festou MC, Kel-
ler HU, Weaver HA (eds) Comets II. University of
The next rendezvous with a comet, provided in Arizona Press, Tucson
2014–2015 by the Rosetta mission, offered Kissel J et al (1986) Composition of comet Halley dust
unique data on the nucleus density and structure, particles from Giotto observations. Nature
as well as on the composition of the dust in the 321:336–337
Korth A et al (1986) Mass spectra of heavy ions near
inner coma and on the nucleus near-surface comet Halley. Nature 321:335–336
(including information on the chirality of the Kr€
uger FR, Korth A, Kissel J (1991) The organic matter of
organic samples). Together with ongoing come- comet Halley as inferred by joint gas phase and solid
tary flybys and remote observations, it is leading phase analyses. Space Sci Rev 56:167–175
Levasseur-Regourd AC et al (1993) Optical probing of
to a better understanding of the suspected link comet Grigg-Skjellerup dust from the Giotto space-
between comets and the origin of life on terres- craft. Planet Space Sci 41:167–169
trial planets. Levasseur-Regourd AC, McBride N, Hadamcik E, Fulle
M (1999) Similarities between in situ measurements of G
local dust scattering and dust flux impact data within
the coma of 1P/Halley. Astron Astrophys
See Also 348:636–641
McBride N, Green S, Levasseur-Regourd AC, Goidet-
Devel B, Renard JB (1997) The inner dust coma of
▶ Comet comet 26P/Grigg-Skjellerup: multiple jets and nucleus
▶ Comet Halley fragments? Mon Not R Astron Soc 289:535–553
▶ Deep Impact Reinhard R (1986) The Giotto encounter with comet Hal-
▶ Deuterium/Hydrogen Ratio ley. Nature 321:313–318
Reinhard R, Battrick B (1986) The Giotto mission, its
▶ Interplanetary Dust Particle scientific investigations. ESA Special Publica-
▶ Rosetta Spacecraft tion1077, Noordwijk
▶ Stardust Mission Rickman H (1989) The nucleus of comet Halley: surface
▶ Vega 1 and 2 Spacecraft structure, mean density, gas and dust production. Adv
Space Res 9(3):59–71
Balsiger H, Altwegg K, Geiss J (1995) D/H and 18O/16O

ratio in the hydronium ion and in neutral water from in
GJ 667C: First System with Multiple
situ ion measurements in comet Halley. J Geophys Res Super-Earths in the Habitable Zone
100:5827–5834
Calder N (1992) Giotto to the comets. Presswork, London Nader Haghighipour
Eberhard P, Reber M, Krankovsky D (1995) The D/H and
18 16
O/ O-ratios in water from comet P/Halley. Astron
Astrophys 302:301–316 Manoa, Honolulu, Hawaii, HI, USA
Fomenkova MN (1999) On the organic refractory
component of cometary dust. Space Sci Rev
90:109–114
Fulle M, Levasseur-Regourd AC, McBride N, Hadamcik
The M1.5V star GJ 667C (Gliese 667C) is a
E (2000) In-situ dust measurements from within the faraway member of a triple star system. It orbits
coma of 1P/Halley: first order approximation with a the central binary GJ 667AB (a K3V+K5V binary
dust dynamical model. Astron J 119:1968–1977 system) at a projection distance of 230 AU. GJ
Grewing M, Praderie F, Reinhard R (eds) (1987) Explora-
tion of Halley’s comet. Springer, Berlin
667C is known to host several super-Earth
Keller HU et al (1986) First Halley multicolour camera planets. The first planet discovered around this
imaging results from Giotto. Nature 321:321–326 star was a hot super-Earth in a 7.2-day orbit
Keller HU, Curdt W, Kramm JR, Thomas N (1994) (Bonfils 2009). However, it was the subsequent
Images of the nucleus of comet Halley obtained by
HMC on board the Giotto spacecraft. In: Reinhard R,
discovery and confirmation of a 4.5 Earth mass
Longdon N, Battrick B (eds) Images of the nucleus of planet, GJ 667Cc, in the habitable zone of this
comet Halley, vol 1. ESA, Noordwijk star (Anglada-Escudé et al. 2012; Delfosse
984 GJ 667C: First System with Multiple Super-Earths in the Habitable Zone
GJ 667C: First System with Multiple Super-Earths in the Habitable Zone, Fig. 1 Artist’s rendering of the triple
star system GJ 667 and planets GJ 667Cb and GJ 667Cc
HZ with HZ with
H2O clouds Habitable Zone CO2 clouds
GJ 667C b h? c f e d g
0.0 0.05 0.10 0.15 0.20 0.25 0.30 0.5

a [AU]
Moist Maximum
greenhouse greenhouse
Recent Early
Venus Mars
0 10 20 30 40 90
d [mas]
GJ 667C: First System with Multiple Super-Earths in the Habitable Zone, Fig. 2 Habitable zone of GJ 667C and
its three potentially habitable super-Earths
et al. 2013) that made this system one of the most PFS/Magellan and HIRES/Keck spectrometers
significant planetary systems known to date. The (Anglada-Escudé et al. 2013) showed that the
planet GJ 667Cc is the first confirmed potentially planetary system of GJ 667C is dynamically
habitable planet. Figure 1 shows an artist’s ren- packed and, in addition to GJ 667Cc, this system
dering of this system. harbors two more super-Earths in its habitable
Reanalyzing the publicly available HARPS zone. Figure 2 shows a schematic of the planets
spectra of GJ 667C using the HARPS-TERRA of this system with respect to its habitable zone.
software (Anglada-Escudé and Butler 2012) As shown here, planets GJ 667Cf and GJ 667Ce,
combined with the Doppler measurements from with orbital periods of 39 and 62 days and masses
Glaciation 985
of 1.94 and 2.68 Earth masses, are also in the characterized by very low surface temperatures
habitable zone. and the advance of glaciers. The term differs from
The discovery of GJ 667Cc as the first con- an “ice age” or more properly a “glacial age” that
firmed potentially habitable planet was consid- is defined as a long-term period of reduction in
ered as one of the top ten physics newsmakers of the temperature of the Earth’s surface and atmo-
2013 by the American Physical Society. sphere, resulting in an expansion of continental
and polar ice sheets.
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been observed on the surface of Mars and have Gliese 581, the Most Highly Debated
been attributed to multiple episodes of glaciation Habitable System
(Kargel and Strom 1992; Dickson et al. 2008).
Indirect evidence of glacial phases on planetary Nader Haghighipour
surfaces could be indirect evidence of chemical Institute for Astronomy, University of Hawaii-
atmosphere evolution and/or active plate Manoa, Honolulu, Hawaii, HI, USA
tectonics.
The planetary system of the M star Gliese 581 is

See Also by far the most highly debated exoplanetary sys-
tem. In 2005, Bonfils et al. announced the detec-
▶ Antarctica tion of a 5.37-day hot Neptune (GJ 581b) planet
▶ Plate Tectonics around this star. Two years after that, a group of
▶ Snowball Earth astronomers from Geneva Observatory published
a paper lead by Stephane Udry in which they
announced the detection of two additional planets
References and Further Reading (Gliese 581c and d) in this system (Udry
et al. 2007). Planet c was reported to have a
Dickson JL, Head JW, Marchant DR (2008) Late Amazo-
nian glaciation at the dichotomy boundary on Mars:
period of 12.931 days and a minimum mass of
evidence for glacial thickness maxima and multiple 5.06 Earth masses. The period and minimum
glacial phases. Geology 36:411–414 mass of planet d were reported to be 83.4 days
Hoffman PF, Kaufman AJ, Halverson GP, Schrag DP and 8.3 Earth masses, respectively. In 2009,
(1998) A neoproterozoic snowball earth. Science
281:1342–1346
Mayor et al. revised the mass and orbital period
Kargel JS, Strom RG (1992) Ancient glaciation on Mars. of planet d to 7.1 Earth masses and 67 days,
Geology 20:3–7 respectively, and announced the detection of a
Gliese 581, the Most Highly Debated Habitable System 987
Gliese 581, the Most Highly Debated Habitable System, Fig. 1 Schematic view of the planetary system of Gliese
581 according to the paper by Udry et al. (2007)
new planet (Gliese 581 e) with an orbital period Gliese 581. However, given a large atmosphere,
of 3.148 days and a minimum mass of 1.9 Earth planet d could harbor surface liquid water (Figs. 2
masses. With these orbital periods, planet c and 3).
would be close to the inner edge of the classical In 2010, Vogt et al. studied worth of 11 years
habitable zone, and planet d would be slightly accumulated HIRES precision radial velocity
outside its outer boundary (Fig. 1). data and showed that the system includes two
Because of the small minimum mass of planet more planets, Gliese 581 f, a 7 Earth-mass planet
c which puts this planet in the regime of poten- in a 433-day orbit, and Gliese 581 g, a 3.1 Earth-
tially rocky super-Earths, its proximity to the mass super-Earth in a ~37-day orbit (Vogt
inner edge of the habitable zone generated con- et al. 2010). Figure 4 shows an artist’s rendering
siderable excitement. An assumed Bond albedo of this system compared to the planets in the solar
of 0.5 yields an equilibrium temperature 320 K system.
for this planet, suggesting the possibility that it The discovery of Gliese 581 g immediately
may be a habitable super-Earth. However, a more caught worldwide attention as, with a semimajor
detailed analysis by Selsis et al. (2007), that axis of ~0.14 AU, this planet is squarely in the
included the greenhouse effect and the spectral habitable zone of its host star (0.1–0.2 AU).
energy distribution of GJ 581, concluded that the Given that this planet is a low-mass super-Earth
surface temperature of planet c is much (Fig. 5), having an orbit in the habitable zone
higher and that this planet is unlikely to host implies that Gliese 581 g could be potentially
liquid water on its surface. Selsis et al. (2007) habitable. And, that way, the detection of this
concluded that both planets c and d are demon- planet marked the discovery of the first habitable
strably outside the conservative habitable zone of world.
988 Gliese 581, the Most Highly Debated Habitable System
Gliese 581, the Most

Highly Debated
Habitable System,
Fig. 2 Schematic view of
the planetary system of
Gliese 581 based on the
paper by Selsis et al. (2007)
Clouds Clouds No clouds CO2 Clouds

100% 50% 100%
Sun Venus Earth Mars
0.0 0.5 1.0 1.5 2.0
Venus Mars
(1 Gyr ago) empirical Habitable Zone (4 Gyrs ago)
G1581 (b) (c) (d)

* * * *
0.00 0.05 0.10 0.15 0.20 0.25
Gliese 581, the Most Highly Debated Habitable System, Fig. 3 Schematic view of the planetary system of Gliese
581 and its habitable zone compared to the solar system (Selsis et al. 2007)
Since the announcement of Gliese 581 g, presented a detailed analysis of the HARPS data
many researchers have examined the available and showed that from that data set alone, one
observational data, in particular the data from cannot detect planets f and g. Unfortunately,
the HARPS spectrograph, and concluded that this paper did not clear the referee stage and has
planet g does not appear in this data set. In remained unpublished.
2011, Forveille et al. submitted a paper to the Subsequent to the analysis by Forveille et al.,
journal Astronomy & Astrophysics where they Vogt et al. (2012) combined the data from
Gliese 581, the Most Highly Debated Habitable System 989
Gliese 581, the Most Highly Debated Habitable System, Fig. 4 Artist’s rendering of the planetary system of
Gliese 581 based on the paper by Vogt et al. (2010)
Gliese 581, the Most Highly Debated Habitable System, Fig. 5 Planets of Gliese 581 as reported by Vogt
et al. (2010)
990 Global Climate Model
HARPS with the data from Keck-HIRES and are known in the ▶ Milky Way, while the giant
reanalyzed the combination of the entire two elliptical galaxy M87 may have up to 10,000. In
data sets. They showed that if circular orbits are the case of the Milky Way, they are composed of
considered for all planets, the two planet f and g old (12 Gyr) and thus low-mass stars, with low
emerge, with planet g having a new orbital period abundances of elements heavier than He, similar
of 32.13 days and revised minimum mass of 2.24 to those of field halo stars. Their high internal
Earth masses and is still in the middle of the density (up to 1,000 stars per pc3 in the cluster
habitable zone. core) favors dynamical interactions between stars
The most recent publication on this system is and makes the survival of any planetary system
by Robertson et al. (2014) who have shown that problematic.
when correcting for stellar activity, the radial
velocity signal of GJ 581 d will diminish imply-
ing that GJ 581 d does not exist but is an artifact
See Also
of stellar activity which, when incompletely
corrected, causes the false detection of planet g.
▶ Galaxy
▶ Milky Way
▶ Star
Bonfils X et al (2005) Astron Astrophys 443:L15
Forveille T et al (2011) ArXiv e-prints 1109.2505, sub-
mitted to Astronomy & Astrophysics
Mayor M et al (2009) Astron Astrophys 507:487
Robertson P, Mahadevan S, Endl M, Roy A (2014) Sci- Globule, Nanoglobule
ence (submitted) (arXiv:1407.1049v1)
Selsis F et al (2007) Astron Astrophys 476:1373 Bradley De Gregorio
Udry S et al (2007) Astron Astrophys 469:L43
Vogt SS et al (2010) Astrophys J 723:954 Materials Science and Technology Division,
Vogt SS, Butler RP, Haghighipour N (2012) U.S. Naval Research Laboratory, Washington,
Astronomische Nachr 333:561 DC, USA
Definition
Global Climate Model
In meteoritic chemistry, globules or
▶ GCM nanoglobules are microscopic, hollow (but some-
times solid), spherical to irregular organic grains
present in the fine-grained matrix of carbona-
ceous chondrites (Fig. 1). Organic globules have
Globular Cluster also been reported in ordinary chondrites, Ant-
arctic micrometeorites, interplanetary dust parti-
Nikos Prantzos cles, and comets. Globule diameters range from
Institut d’Astrophysique de Paris, Paris, France less than 100 up to 2 mm. Many are enriched in
▶ deuterium and/or nitrogen-15 relative to the
surrounding bulk ▶ insoluble organic matter,
Definition suggesting formation in extremely cold (less
than 20 K) ▶ molecular cloud or ▶ solar nebula
A globular cluster is a dense spherical collection environments where such isotopic enrichments of
of 105–106 stars orbiting a galaxy. Their origin is organic matter are theorized to occur (Nakamura-
poorly understood. About 160 globular clusters Messenger et al. 2006). Some organic globules
Glucogenesis 991
Claus G, Nagy B (1961) A microbiological examination

of some carbonaceous chondrites. Nature
192:594–596
Cody GD et al (2011) Establishing a molecular relation-
ship between chondritic and cometary organic solids.
Proc Natl Acad Sci USA 108:19171–19176
Garvie LAJ, Buseck PR (2004) Nanosized carbon-rich
grains in carbonaceous chondrite meteorites. Earth
Nakamura K et al (2002) Hollow organic globules in
the Tagish Lake meteorite as possible products
of primitive organic reactions. Int J Astrobiol
1:179–189
Nakamura-Messenger K et al (2006) Organic globules in
the Tagish Lake meteorite: remnants of the protosolar G
disk. Science 314:1439–1442
Globule, Nanoglobule, Fig. 1 Transmission electron

microscopy image of an organic nanoglobule within
fine-grained silicate matrix of ordinary chondrite QUE
97008 Glove Box
Ricardo Amils
may also have formed on asteroid and/or plane- Departamento de Biologia Molecular,
tesimal parent bodies through organic polymeri- Universidad Autónoma de Madrid, Madrid, Spain
zation, such as the ▶ formose reaction, during
aqueous processing (Cody et al. 2011).
Definition
History A glove box is a sealed container that is designed

to manipulate objects in a different atmosphere.
Originally discovered in 1961 in crushed Built into the front of the glovebox are gloves
samples of the Orgueil and Ivuna meteorites by arranged in such a way that the user can place
George Claus and Bartholomew Nagy (Claus their hands into the gloves and perform tasks
and Nagy 1961), organic globules were inside the box without breaking containment.
initially interpreted as ▶ microfossils indigenous Two types of glove boxes exist: one manipulates
to their host meteorites. They were hazardous substances, such as infectious disease
“rediscovered” independently by Keiko agents; the other allows manipulation of sub-
Nakamura-Messenger in 2002 (Nakamura stances in an oxygen-free or a high-purity inert
et al. 2002) and Lawrence Garvie in 2004 atmosphere. It is very useful for the manipulation
(Garvie and Buseck 2004). of strict anaerobic microorganisms.
See Also
▶ Carbonaceous Chondrites, Organic Chemistry of Glucogenesis

▶ Insoluble Organic Matter
▶ Nitrogen Isotopes ▶ Gluconeogenesis
992 Gluconeogenesis
Glutamic Acid,
Gluconeogenesis Fig. 1 Glutamic acid
Juli Peretó
Spain
Synonyms
Glucogenesis
Glutamic Acid,
Fig. 2 Pyroglutamic acid
O COOH
N
Definition H
Gluconeogenesis refers to the biosynthetic pathway

of hexoses from noncarbohydrate substrates such as and one-letter symbol is E. Its compositional
acetate, glycerol, pyruvate, etc. This can be structure is C5H9N4, and its molecular weight is
achieved by conversion of the precursor into an 147.13. It is a monoaminodicarboxylic acid and is
intermediate of ▶ glycolysis and the reverse of classified as an acidic ▶ amino acid, whose iso-
this pathway (with the appropriate bypass of three electric point (pI) is 3.22. Its salt is referred to as
irreversible enzymatic steps). The phylogenetic dis- glutamate: monosodium glutamate is a popular
tribution of the gluconeogenetic enzymes suggests condiment. When it is heated, intramolecular
an early origin of this anabolic pathway, before the condensation occurs to give pyroglutamic acid
emergence of the different glycolytic pathways. (Fig. 2). It is found in extracts from carbonaceous
chondrites and is easily produced in a number of
See Also prebiotic synthesis experiments such as electric
discharge experiments.
▶ Anabolism
▶ Embden-Meyerhof-Parnas Pathway See Also
▶ Entner-Doudoroff Pathway
▶ Glycolysis ▶ Amino Acid
▶ Metabolism ▶ Protein
Glutamic Acid Glutamine
Kensei Kobayashi Kensei Kobayashi

Yokohama National University, Tokiwadai, Yokohama National University, Tokiwadai,
Hodogaya-ku, Yokohama, Japan Hodogaya-ku, Yokohama, Japan
Glutamic acid (Fig. 1) is one of the 20 ▶ protein Glutamine is one of the 20 protein ▶ amino acids,
amino acids, whose three-letter symbol is Glu whose three-letter symbol is Gln and one-letter
Glycerol 993
symbol is Q. Its side chain has a terminal amide basic aqueous solution. It has not been detected in
group (-CONH2). The chemical formula for glu- carbonaceous chondrites nor in interstellar space,
tamine is C5H10N2O3, which gives a molecular although the related triose, dihydroxyacetone
weight of 146.14. It has an isoelectric point (pI) (CH2OH-CO-CH2OH), has been reported in
of 5.65. It is easily hydrolyzed to give the amino both. However these interstellar identifications
acid ▶ glutamic acid (Glu) and ammonia. Both have recently been questioned.
glutamic acid and glutamine are determined as
glutamic acid in the hydrolysate of ▶ proteins.
After acid hydrolysis, it is thus unknown whether
See Also
the original species was Glu or Gln, and thus it is
reported as Glx.
▶ Carbohydrate
▶ Formaldehyde
G
See Also ▶ Homochirality
▶ Amino Acid
▶ Glutamic Acid
▶ Protein
Glycerin
▶ Glycerol
Glyceraldehyde
Kensei Kobayashi
Glycerine
Hodogaya-ku, Yokohama, Japan
▶ Glycerol
Definition
Glyceraldehyde is a one of the simplest ▶ carbo- Glycerol

hydrates (also called sugars); its chemical struc-
ture is CH2OH-CH2OH-CHO. It is an aldo-triose Henderson James (Jim) Cleaves II
(three carbon sugar with an aldehyde group). Earth–Life Science Institute (ELSI), Tokyo
Since the middle carbon atom is stereogenic Institute of Technology, Meguro–ku, Tokyo,
(connected to four different groups: H, OH, Japan
CHO, and CH2OH), it is a chiral compound and Institute for Advanced Study, Princeton,
two enantiomeric forms are possible: NJ, USA
D-glyceraldehyde and L-glyceraldehyde. Terres- Blue Marble Space Institute of Science,
trial organisms largely use only Washington, DC, USA
D-glyceraldehyde. D-Glyceraldehyde (as its Center for Chemical Evolution, Georgia Institute
phosphate ester) plays important roles in sugar of Technology, Atlanta, GA, USA
metabolism, such as in the Benson-Calvin cycle
of photosynthesis and in glycolysis. It can be
synthesized by mild oxidation of ▶ glycerol. It Synonyms
is formed in the formose reaction, i.e., the poly-
merization of ▶ formaldehyde with a catalyst in 1, 2, 3-Propantriol; Glycerin; Glycerine
994 Glycine
Definition was proven by Miller-Urey type experiments

using various types of primitive atmospheric gas
Glycerol is a three-carbon organic compound with mixtures and energy sources.
a hydroxyl group on each carbon atom. It is vis-
cous colorless liquid at room temperature. It is
Overview
extremely soluble in water and is a component of
many lipids, either as part of fatty acid esters or
▶ Glycine is the simplest amino acid and is
phytanyl ethers. Dissolved aqueous glycerol
formed readily and apparently ubiquitously
lowers the freezing point of water. Glycerol, as
under various natural conditions. Its pKa values
its phosphate ester, is generally metabolically
are 2.35 and 9.78 at 25 C; in neutral solution, it
derived from the reduction of glyceraldehyde 3-
exists therefore predominantly as its ▶ zwitter-
phosphate or dihydroxyacetone phosphate.
ion, +H3NCH2CO2 .
Glycine has been identified in meteorites and
comets (Elsila et al. 2009). Although the detec-
Glycine tion of interstellar glycine remains controversial
(see ▶ Molecules in Space), the glycine precursor
Bernd Michael Rode aminoacetonitrile was discovered in SgrB2, a
Institute for General, Inorganic and Theoretical giant gas cloud near the galactic center
Chemistry, Leopold-Franzens University, (Belloche et al. 2008). Glycine could also have
Universität Innsbruck, Innsbruck, Austria formed easily from reactions in the primitive
atmosphere under various conditions, such as
the classical Miller-Urey setting via electrical
Keywords discharges acting on an atmosphere of hydrogen,
methane, ammonia, and water (Miller 1953), or a
Amino acids; Chemical evolution; Glycine; Ori- carbon dioxide – N2 – water atmosphere
gin of life; Peptides; Proteins (Plankensteiner et al. 2004), now assumed to
have predominated on Earth 3.8 billion years ago.
A number of other experiments have also demon-
Synonyms strated the formation of glycine from basic
chemicals under the influence of various energy
Aminoacetic acid; Aminoethanoic acid; sources. It can be derived via Strecker synthesis
Glycocoll; NH2CH2COOH from NH4CN and formaldehyde, as well as directly
from the self-condensation of HCN.
Starting from glycine, numerous plausibly
Definition
prebiotic chemical reactions lead to more com-
plex amino acids which are also important in
Glycine is the simplest ▶ amino acid. It is achiral
biochemistry, such as serine and alanine
(see ▶ Chirality) and found in ▶ meteorites and
(Choughuley et al. 1975). Hence, glycine may
▶ comets, indicating the ease of its abiotic for-
have enabled the formation of complex peptides
mation. It is catalytic in some peptide formation
even before the advent of RNA.
reactions and can be readily oligomerized under
some geochemically plausible conditions.
See Also
History ▶ Amino Acid

▶ Carbonaceous Chondrites, Organic Chemistry of
Glycine was first isolated by Henri Braconnot in ▶ Protein
1820. Its formation under prebiotic conditions ▶ Strecker Synthesis
Glycolaldehyde 995

Glycolaldehyde
Belloche A, Menten KM, Comito C, M€ uller HSP,
Schilke P, Ott J, Thorwirth S, Hieret C (2008) Detec-
Didier Despois
tion of amino acetonitrile in Sgr B2(N). Astron
Astrophys 482:179–196 Laboratoire d’Astrophysique de Bordeaux,
Bernd M (2007) The first steps of chemical evolution CNRS-Universite de Bordeaux, France
towards the origin of life. Chem Biodivers
4:2674–2702
Choughuley AS, Subbaraman AS, Kazi ZA,
Chadha MS (1975) Transformation of some hydroxy Synonyms
amino acids to other amino acids. Orig Life
6(4):527–535 HOCH2CHO; Hydroxyacetaldehyde
Elsila JE, Glavin DP, Dworkin JP (2009) Cometary gly-
cine detected in samples returned by stardust. Meteorit
Planet Sci 44(9):1323–1330
Miller SL (1953) A production of amino acids under Definition G
possible primitive earth conditions. Science
117(3046):528–529
Glycolaldehyde (IUPAC name hydroxyace-
Plankensteiner K, Reiner H, Schranz B, Rode BM
(2004) Prebiotic formation of amino acids in a neutral taldehyde) is the simplest molecule that is both
atmosphere by electric discharge. Angew Chem Int Ed an alcohol and an aldehyde. It shares with
43:1886–1888 ▶ monosaccharides (“sugars”) a formula
Rode BM (1999) Peptides and the origin of life. Peptides
(H2CO)n so that it can be considered formally to
20(6):773–786
be a polymer of ▶ formaldehyde H2CO and to
be a carbohydrate. It is an intermediate in
the ▶ formose reaction. Monosaccharides have
n 3, so that glycolaldehyde is a “pre-sugar”
Glycine Anhydride rather than a sugar. Glycolaldehyde has been
identified in the interstellar medium.
▶ Diketopiperazine
History
Hollis et al. (2000) identified glycolaldehyde for

the first time in the interstellar medium, toward
Glycinonitrile
the Galactic center.
▶ Aminoacetonitrile
See Also
▶ Carbohydrate
Glycocoll ▶ Formaldehyde
▶ Glycine ▶ Molecules in Space
▶ Monosaccharide

Glycol Hollis JM, Lovas FJ, Jewell PR (2000) Interstellar
glycolaldehyde: the first sugar. Astrophys J 540:
▶ Ethylene Glycol 107–110
996 Glycolic Acid
electrons. In anaerobic conditions, glycolysis can

Glycolic Acid be the only source of ▶ ATP through the
substrate-level phosphorylation mechanism.
Henderson James (Jim) Cleaves II The diversity of glycolytic pathways
Earth–Life Science Institute (ELSI), Tokyo includes the ▶ Embden-Meyerhof-Parnas path-
Institute of Technology, Meguro–ku, Tokyo, Japan way (the canonical version of glycolysis) present
Institute for Advanced Study, Princeton, NJ, USA in bacteria and eukaryotes, the variants of the
Blue Marble Space Institute of Science, ▶ Entner-Doudoroff pathway found in
Washington, DC, USA bacteria and archaea, and the hexose
Center for Chemical Evolution, Georgia Institute monophosphate pathway (or pentose phosphate
of Technology, Atlanta, GA, USA pathway). Different redox balance mechanisms
under anaerobic conditions allow the operation
Synonyms of a diversity of ▶ fermentation pathways in
microorganisms.
2-Hydroxyethanoic acid
See Also
Definition
▶ Embden-Meyerhof-Parnas Pathway
Glycolic acid, HOCH2COOH, is the smallest ▶ Entner-Doudoroff Pathway
a-hydroxy acid. It is an odorless and colorless ▶ Fermentation
hygroscopic crystalline solid at room tempera- ▶ Metabolism
ture, and it is highly soluble in water. It plays a
minor role in biochemistry and has been found in
carbonaceous chondrites and in electric discharge
experiments. The hydrolysis of cyanohydrin Glycolyurea
formed from HCN and formaldehyde yields
glycolamide and finally glycolic acid, which ▶ Hydantoin
may be the source of this compound in meteorites
and certain types of prebiotic syntheses.
See Also
Goethite
Daniele L. Pinti
▶ Hydroxy Acid
Glycolysis
Synonyms
Juli Peretó
Acicular ironstone; Limonite needle ironstone
Glycolysis refers to the metabolic pathways using Goethite is a hydrated iron oxide, or ▶ iron
carbohydrates as a source of carbon, energy, and oxyhydroxide, of chemical formula Fe3+O
Gondwana 997
(OH) (orthorhombic crystal system). Goethite He systematized the use of X-rays in study of
forms by ▶ weathering of iron-rich minerals and composition and structure of crystals.
it is commonly found in soils. It may also be Goldschmidt wrote a manuscript about geochem-
precipitated by groundwater in low-temperature ical aspects of the origin of complex organic mol-
environments (authigenic goethite in marine and ecules on the Earth.
lacustrine sediments). Silica-associated goethite
has been found in hydrothermal deposits. The
See Also
occurrence of goethite on planetary surfaces
could be used as an indicator of
▶ Chalcophile Elements
low-temperature ▶ weathering with the involve-
▶ Lithophile Elements
ment of liquid water.
▶ Rare Earth Elements
▶ Siderophile Elements
G
See Also

Gondwana
▶ Hematite
▶ Iron
Nicholas Arndt
▶ Iron Cycle
▶ Iron Oxides, Hydroxides and Oxy-hydroxides
▶ Mars
▶ Weathering
Synonyms
Gondwanaland
Goldschmidt, Viktor Moritz

Definition
Pierre Savaton
Université de Caen Basse-Normandie, Caen, Gondwana (originally Gondwanaland) is the
France ▶ supercontinent composed of the land masses
of the present-day southern hemisphere (Africa,
South America, Australia, and Antarctica,
History together with India and Arabia). Together with
its northern counterpart, ▶ Laurasia, Gondwana
Victor Moritz Goldschmidt (1888–1947) is con- comprised almost all of present-day continental
sidered the founder of modern geochemistry, crust. Between about 500 and 200 Ma, Gondwana
establishing its general laws and principles and and Laurasia together formed part of a single
making it a new discipline in earth sciences. supercontinent, ▶ Pangea. Breakup started
Among his works, there are researches about min- 180–200 Ma ago when Gondwana split from
eral metamorphic associations and thermody- Laurasia. It accelerated about 170 Ma ago when
namic equilibrium; the laws which determine Antarctica, Madagascar, India, and Australia
distribution of the chemical elements on the began to separate from Africa, and the Atlantic
Earth and their classification as litho- (rock- Ocean progressively opened to separate South
loving), sidero- (iron-loving), and chalcophile America from Africa. The correlation of geolog-
(sulfur-loving); the ionic substitution in mineral; ical features and fossils on both sides of the
the process of fractional crystallization; and Atlantic provided convincing evidence of conti-
the substitution laws of ions of rare elements. nental drift and eventually ▶ plate tectonics.
998 Gondwanaland
History ▶ peptidoglycan and an ▶ outer membrane

containing lipopolysaccharides, proteins, and
Austrian geologist Eduard Suess named other complex macromolecules. The distinction
Gondwana after a region in northern India (from between Gram-positive and ▶ Gram-negative
Sanskrit gondavana which means “forest of bacteria is based on a differential stain, the
Gond”). Gram stain, which depends on the structure of
the cell wall (see cell wall). After Gram staining,
Gram-positive bacteria appear purple and Gram-
negative bacteria appear red. This difference in
See Also reaction to the Gram stain arises because of dif-
ferences in the cell wall structure which promote
▶ Laurasia the retention of the purple dye, crystal violet, in
▶ Pangea the Gram-positive bacteria after decoloring with
▶ Plate Tectonics ethanol, while in the Gram-negative bacteria, the
▶ Supercontinent stain is not retained. The Gram stain is one of the
most useful staining procedures in microbiology.
In addition to the peptidoglycan, Gram-negative
bacteria contain an additional wall layer, the
outer membrane. This layer is effectively a sec-
ond lipid bilayer, but in addition to phospholipids
Gondwanaland and proteins as in the cytoplasmic membrane, it
contains polysaccharides. The lipid and polysac-
▶ Gondwana charide are linked to form a lipopolysaccharide
complex known as LPS. Unlike the cytoplasmic
membrane, the outer membrane of Gram-
negative bacteria is relatively permeable to
small molecules due to the presence of porins,
Graben proteins that function as channels that facilitate
the transit of low-molecular-weight substances.
▶ Rille The outer membrane is not permeable to proteins
▶ Rima, Rimae and other large molecules. The space between the
▶ Vallis, Valles outer and the cytoplasmic membranes is known
as ▶ periplasm. In the periplasm, important func-
tional proteins like hydrolytic enzymes or bind-
ing proteins for transport acting as
chemoreceptors can be found.
Gram-Negative Bacteria
Ricardo Amils
Departamento de Biologia Molecular, See Also
▶ Bacteria
▶ Cell Wall
▶ Chemotaxis
Definition ▶ Cytoplasmic Membrane
▶ Outer Membrane
A Gram-negative bacterium is a prokaryotic cell ▶ Peptidoglycan
with a ▶ cell wall containing small amounts of ▶ Periplasm
Granite 999
▶ Cell Wall
Gram-Positive Bacteria ▶ Gram-Negative Bacteria
▶ Peptidoglycan
Ricardo Amils ▶ Phylogeny
Granite
Definition
Nicholas Arndt
A Gram-positive bacterium is a prokaryotic cell
whose ▶ cell wall consists of mainly ▶ peptido-
glycan and lacks the outer membrane characteristic
of the Gram-negative cells. Bacteria can be divided G
into two major groups, called Gram positive and Definition
Gram negative. The original distinction between
both groups was based on the Gram stain, which Granite is a coarse-grained igneous intrusive
depends on differences in the structure of the cell rock composed mainly of Si- and Al-rich min-
wall (see ▶ Cell Wall). The Gram-negative cell wall erals (quartz and feldspars), with minor amounts
is a mutilayered structure and quite complex, of biotite, muscovite, and amphibole. Granitic
whereas the Gram-positive cell wall consists of a rocks are the dominant component of the ▶ con-
single type of molecule, the peptidoglycan, which tinental crust. A granite cannot be a primary melt
is often much thicker. In Gram-positive bacteria, as of mantle peridotite. Its genesis (and thus that of
much as 90 % of the cell wall consists of peptido- continental crust) involves the fractional crystal-
glycan, although another molecule, teichoic acid, is lization of mafic magmas or more commonly,
usually present in small amounts. The phylogenetic remelting of basaltic or sedimentary precursors.
analysis of bacteria has demonstrated that the cell Granite has not been reported on planets other
wall structure is phylogenetically consistent: all than Earth, except as rare disseminated veinlets in
Gram-positive bacteria form a coherent phylogenetic a few lunar rocks. Most granite forms in a
group. The Gram-positive bacteria are a large group two-stage process: The first stage involves partial
of chemoorganotrophic bacteria. The Gram-positive melting of the lower crust and sometimes of
bacteria can be separated into two subgroups, subducted oceanic material (of basaltic composi-
according to the GC content of their genome. tion) to form magmas of intermediate andesitic
Among the different Gram-positive, low-GC bacte- composition. Andesite, in turn, partially melts to
ria of industrial interest can be found, including lactic form magmas of granitic composition. S-granites
acid bacteria, endospore-forming bacteria (like those are intrusive rocks formed by the partial melting
belonging to the Bacillus and Clostridium genus), of metasedimentary and metavolcanic rocks
and cell-wall-less bacteria, like the Mycoplasma. within orogenic belts through the process of
The high-GC, Gram-positive bacteria group corre- anatexis.
sponds to the phylum Actinobacteria, some of them
producers of antibiotics, secondary metabolites of
pharmacologic interest.
See Also
See Also ▶ Continental Crust

▶ Crust
▶ Antibiotic ▶ Igneous Rock
▶ Bacteria ▶ Mafic and Felsic
1000 Graphite
Synonyms
Graphite
Gravity
Francis Albarède
France Definition
Definition Gravitation is one of the four fundamental forces

of physics. It causes an attraction between any
Graphite is a mineral composed of carbon arranged two massive bodies which is proportional to their
in graphene sheets of six-atom hexagonal rings and masses.
stacked in layers. It is the stable form of crystalline
▶ carbon at the Earth’s surface and shallow pressure
and with increasing pressure converts to diamond, Overview
which in the Earth is stable at depths in excess of
150 km. Graphite is a widespread accessory mineral Gravitation is a property of space which modifies
in both terrestrial rocks and meteorites. It may result the velocity of massive bodies under its influence.
from the inorganic reduction of carbon dioxide and Then, by using Newton’s definition, it is a force.
carbonate, notably by hydrogen, or from the dehy- In the context of classical (i.e., nonrelativistic)
drogenation of organic materials at temperatures in physics, this same Newton gave a formula for
excess of 300 C. Typically, it crystallizes during gravity as a fundamental example of his theoret-
metamorphism from precursor carbonaceous mate- ical work between 1666 and 1687. This is the
rial or forms from fluid deposition as a precipitate of famous inverse-square law: the force produced
CO2CH4H2O. It coexists with methane-rich by a point-like body of mass mA located at the
gases over a broad range of temperatures. The disposition A on another point mass mB located at the
covery of 13C-depleted graphite in 3.8 Ga-old rocks position B is given by Fig. 1.
from West Greenland triggered a controversy over The second formulation, in which the squared
whether they represent carbon from early organic distance d2 appears, is the more popular but needs
!
remains or inorganic precipitates. to introduce the unit vector u in the direction
from one mass toward the other.
See Also If an apple falls, its velocity changes: from
zero when it leaves the tree to a certain value
▶ Carbon when it reaches the ground. This apple is thus
▶ Carbon Isotopes as a Geochemical Tracer subject to a force.
▶ Kerogen If the Moon orbits around the Earth, it is also
because its velocity is slightly modified at each
instant, by changing its direction in order to close
its orbit. As the apple, the Moon is thus subject to
Gravitation a force. Newton had the idea to consider that
these two forces are of the same nature: they are
Jérôme Perez of gravitational origin.
Applied Mathematics Laboratory, ENSTA On one hand, situated on the surface of the
ParisTech, Paris, Cedex 15, France Earth and at one Earth radius (6,400 km) from
its center, the apple falls 5 m in 1 s. On the other
hand, each second, the orbit of the Moon moves
Keywords away from a straight line by 1.4 mm. Finally,
Newton knew that the Moon was located at a
Fundamental interaction; Mass distance representing 60 times the radius of the
Gravitation 1001
u
B
AB
F A→B = –G mA mB d mB
3
⏐AB⏐
F A →B
G mA mB mA
=– u
d2 A
Gravitation, Fig. 1 Definition of the geometrical quantities in the Newtonian gravitational force
Earth. The apple, which is then nearer to the Earth constants and is independent of the distance. Its
by this same factor, feels an effect which is magnitude, of order 1040 in favor of electrostat-
5/0.0014 602 stronger, hence the inverse-square ics, makes gravity irrelevant for most chemical
law! The other quantities which appear in the problems. Nevertheless, as masses are always of G
formula are the respective masses of the the same sign, gravity is always attractive,
interacting bodies and Newton’s universal gravita- whereas the electrostatic force can be repulsive.
tional constant G, needed to fix the intensity of It must be noted at this point that in general
the force. Newton didn’t precisely know the relativity, repulsive gravity can appear; it is
value of this constant; its measure requires precise caused by the so-called dark energy and could
and refined experiments. One of the first values, be relevant for dynamics at a very large scale,
G ¼ 6:7 1011 m3 kg1 s2, was obtained by using such as in galaxy clusters or in the context of the
a torsion balance and was proposed by Cavendish primordial universe.
in 1798. This is why some texts refer to This purely attractive behavior of the classical
Cavendish’s constant. The modern value, gravitational force represents the main character-
G ¼ ð6:67428 0:00067Þ 1011 m3 kg1 s2 , was istic of gravity; although it is the weakest funda-
recommended in 2006 by the CODATA (see Mohr mental interaction, it cannot be screened. This
et al. 2006). feature allows gravity to build and shape objects
The more complete description of gravitation on very large spatial and temporal scales.
is Einstein’s theory of general relativity. This Pure gravitational problems are generally
formulation confirms the deep relation between classified according to the number of interacting
gravitation and space, including its temporal bodies. On one hand, two-body problems and
dimension, which is absent in the Newtonian their perturbations are the domain of celestial
theory. The refinements introduced by Einstein mechanics. On the other hand, when the number
are fundamental for theoretical physics, but in of interacting bodies is more than, say, several
most astrophysical cases, the Newtonian formu- thousands, like in star clusters or cluster of gal-
lation of gravity allows a correct description of axies, one deals with a sort of self-gravitating gas
the general properties of the system (the Newto- and thermodynamics law may be used. There also
nian theory breaks down, e.g., in the vicinity of a exist systems for which gravity is only one among
black hole). the fundamental interactions that must be consid-
The form of the gravitational force equation is ered. That is especially the case for stars or large
equivalent to the electrostatic one (Coulomb’s gaseous planets on which gravity acts to keep the
law), but in the latter case, the masses matter confined, whereas internal pressure pushes
are replaced by electric charges and Newton’s it outward. The balance between these opposite
constant is replaced by (4pe0)1 where e0 = forces produces, at equilibrium, a star or a planet.
8.854 1012A2s4kg1m3. Due to their same The different states in which we observe stars
spatial dependency, the ratio of the electrostatic correspond to the different types of pressure
force to the gravitational one between protons (thermodynamical, degeneracy) that must be
and electrons depends only on fundamental considered to balance the gravity associated to
1002 Gravitation
their respective masses. The isotropic character trajectories are quite exceptional; ellipsoidal is
of gravity is at the basis of the spherical aspect of the rule. In the solar system, all the planets lie
large gaseous planets or stars. In the case of on such orbits, as do satellites around planets.
telluric planets, the situation is more compli- This configuration provides an equilibrium state
cated: gravity produces a sphere as soon as the between the central force imposed by gravity and
body’s radius exceeds some hundreds of the centrifugal one due to this kind of orbit. This
kilometers. last remark explains why the velocity of B is not
uniform: Near the focus (planet at perihelion for
The Two-Body Problem and Its Perturbations motion around the Sun), gravity is strong and
The academic two-body problem deals with two must be balanced by a large velocity; on the
point masses A and B evolving in empty space contrary, at aphelion, the velocity reaches a min-
and interacting only through their reciprocal imum. Such elliptic orbits are periodic with a
1=2
Newton force (see Fig. 1). This problem was period T ¼ 2pða3 =mÞ where a is the semimajor
solved by Newton in 1687. The defining equation axis of the ellipse. This relation is the well-known
is Kepler’s third law.
When B is not a point mass and if its extension
! !
d2 r r is large enough with respect to the gradient of the
¼ m 3 gravity field, tidal effects can occur. The princi-
dt2 !
r pal tidal effect is an elongation produced by the
stretching (see the entry ▶ Roche limit). This
!
where r is the vector distance between the elongation, combined with B’s own spin at an
masses (the distance AB in Fig. 1) and m depends angular velocity o, produces a torque (in green
on the chosen reference frame. In celestial on Fig. 2b). This torque accelerates or decelerates
mechanics, one of the two bodies (say A) is gen- the spin of B until o reaches the value of B’s
erally much heavier than the other. If we consider angular velocity of revolution O around A.
the reference frame in which A is at rest, one has This is the reason why the Moon always shows
! ! !
r ¼AB and m ¼ GðmA þ mB Þ. The solution r ðtÞ the same face to the Earth: Because of the tidal
of this problem is a conic curve, the nature of effect from the Earth, the Moon takes the same
which depends on B’s total energy EB. This time to rotate around its axis as it does to revolve
energy is the sum of B’s kinetic energy which is around the Earth. This phenomenon is called
positive and of B’s potential gravitational energy synchronization. The reciprocal effect (from
which is by definition negative. If EB 0, the Moon to Earth) makes Earth rotate more slowly
conic is a hyperbola or a parabola with focus on itself: The length of the day actually increases
A and the curve is open. If EB < 0, the conic is by 2 m per century. For instance, at the end of the
an ellipse with A at one focus (this case includes Neoproterozoic (900 million years ago), each
the circle), and the curve is closed. Open year on Earth was lasting 481 days, 18 h. Tidal
a Elongation b Synchronisation
Direction
of A ω
Ω
B B
Gravitation, Fig. 2 Cartoon describing the two major tidal effects

Gravitational Biology 1003
Rotation of the
orbital plane
A Precession of pericenter
in the orbital plane
A
Ref Movement of the
ere
pla nce line of nodes
ne
Gravitation, Fig. 3 Principal effects of a radial perturbation on the elliptical two-body problem G
effects can also circularize the orbit of planets or ▶ Roche Limit

satellites that were initially in an eccentric orbit. ▶ Surface Gravity
This last phenomenon is generally less effective
than the previous one and only takes place for
planets or satellites in the inner solar system. It
does seem to occur for short-period exoplanets. Landau L, Lifshitz E (1976) Mechanics – course of theo-
Perturbations of the two-body problem occur retical physics – Tome 1, 3rd edn. Pergamon, Oxford
when the source of the potential (essentially A) Misner CW, Thorne KS, Wheeler JA (eds) (1973) Gravi-
cannot be considered as a point mass or when tation. W. H. Freeman, San Francisco
Mohr PJ, Taylor BN, Newell DB (2006) CODATA
other bodies must be considered. Generally, recommended values of the fundamental physical con-
small perturbations produce a precession of the stants. Rev Mod Phys 80:633–730
perihelion and a rotation of the orbital plane. The Murray CD, Dermott SF (1999) Solar system dynamics.
intersection, called line of nodes, between a ref- Cambridge University Press, Cambridge
Perez J (2008) Gravitation classique: Problème à N corps,
erence plane (the ecliptic for planets moving de 2 à l’infini, Les presses de l’ENSTA
around the Sun) and the orbital plane then gener-
ally rotates (Fig. 3).
For example, the Sun produces a precession of
the Moon’s perigee (closest approach to Earth) Gravitational Biology
and a regression of this orbit’s line of nodes
around the Earth. These two motions have the Ruth Hemmersbach1, Ralf H. Anken1 and
same period of about 18 years. Most of the Michael Lebert2
1
motions in solar or stellar systems can be German Aerospace Center (DLR), Institute of
explained using Newton’s theory coupled with Aerospace Medicine, Cologne, Germany
2
perturbations and tidal effects. The only “remark- Biology Department, Plant Ecophysiology,
able” exception is the 42 arcsec per century Friedrich-Alexander-University Erlangen/
motion of Mercury’s perihelion, which has been Nuremberg, Erlangen, Germany
explained by Einstein’s theory of gravity.
Keywords
See Also
Behavior; Centrifuges; Clinostat; Development;
▶ Keplerian Orbits Evolution; Gravisensors; Gravitaxis;
▶ Orbital Resonance Gravitropism; Gravity; Hypergravity;
1004 Gravitational Biology
Microgravity; Signal transduction chains; Stato- Basic Methodology), obvious effects such as
cyst; Statocyte; Statolith; Weightlessness gravitaxis (orientation of free-moving organisms
with respect to gravity), gravitropism (orientation
and growth direction of sessile organisms), and
Definition general physiological changes (see Key Research
Findings) have been identified. However, much
The main scope of gravitational biology is the remains to be learned about the impact of gravity
identification and understanding of the effects of on basic mechanisms and about possible counter-
gravity on organisms. Main emphases are the measures to effects which occur during the
identification of gravity sensing systems as well absence of gravity/under ▶ microgravity
as the analysis of gravity-sensitive responses on conditions.
all biological organizational levels. This includes
the clarification of the role of gravity during
evolution and also during individual develop- Basic Methodology
ment, in the course of which gravity has shaped
life and provides an essential cue for spatial Ground-based studies under conditions of
orientation. hypergravity (by the use of centrifuges) or artifi-
cial (simulated) weightlessness (by the use of 2-D
and 3-D clinostats/random positioning machines
History and magnetic levitation) have identified and
brought deeper insights into gravity-induced
The origin of the discipline “Gravitational biological phenomena. Subsequently, such
Biology” might be set at the nineteenth century, original studies are being verified in real micro-
when Julius Sachs (1868), Charles Darwin gravity (free fall, near weightlessness) with time
(1880), and Wilhelm Pfeffer (1904) started to spans ranging from seconds to several minutes,
investigate the influence of gravity on plants by which can be accomplished using, e.g., drop
demonstrating the role of the root cap in down- tower, parabolic aircraft, and ballistic rocket
ward growing plants (gravitropism). Sachs and flights. Long-term studies (weeks to months)
Pfeffer already constructed machines require the use of orbiting spacecrafts such as
(centrifuges and clinostats, see Basic Methodol- satellites or space stations (e.g., MIR, ISS). The
ogy) in order to change the influence of gravity methodology for analyses depends on the issue
and observed the resulting gravity dependent under investigation and ranges from qualitative
response. video observation to quantitative molecular biol-
ogy and high-throughput technologies (-omics).
Control experiments are designed to carefully
Overview distinguish between direct effects of gravity ver-
sus indirect effects due to changing environmen-
Gravity is one of the most important environmental parameters, such as gravity dependent
tal factors which organisms use for orientation alterations in convection or general stress
and adaptation to their environment to optimize responses.
exploitation of resources. In contrast to all other
environmental factors such as light/radiation,
temperature and humidity, gravity has always Key Research Findings
been constant in direction and strength during
evolution of life. Gravity sensing mechanisms Studies in order to identify the influence of grav-
have evolved early in the phylogenesis of organ- ity on living material cover all organizational
isms. By studying the influence of altered gravity levels including isolated proteins, single cells,
by increasing or decreasing Earth gravity (see tissues, and whole organisms.
Cellular and Molecular Biology cytoskeleton-related organization processes

The observation that already unicellular organ- (including the role of the cytoskeleton in
isms (protists, protozoa) can swim to the top of a graviresponding) as well as concerning gravity-
completely filled and sealed tube by means of related genes and the signal transduction of gravity
gravitaxis has been made more than a century signaling. However, the molecular mechanisms of
ago. A real breakthrough in understanding the graviperception in cells remain widely unknown.
underlying mechanisms came with technologies
designed in order to alter the influence of gravity Animal Biology
and methodological advances such as image anal- Spatial orientation and development of animals
ysis of cellular behavior as well as biochemical requires the complex processing of a variety of
approaches. Due to current knowledge, stimulus inputs by means of specific receptors.
graviperception in the ciliate Paramecium and Thus it seems likely that altered gravity impairs
the flagellate Euglena is based on mechano- the physiology of organisms on numerous levels.
(gravi-)sensitive ion channels in the cell mem- Microgravity affects early developmental pro- G
brane which are stimulated by the mechanical cesses; however, regulatory capabilities are able
load of the mass of the cytoplasm. Very few to correct and compensate at least some of these
exceptional species (Loxodes, ciliate) have modifications via more or less unknown mecha-
already developed statocyst-like organelles for nisms, resulting in an apparently “normal” adult
perceiving the direction of gravity. Recent cell animal compared to the Earth-grown control.
and molecular biological studies in microgravity Experiments employing aquatic vertebrates
and hypergravity concentrated on the cytoskele- yielded critical periods for the development of
ton and signal transduction chains, as they control the vestibular system. Multiple adaptive effects
fundamental cellular functions, such as mainte- of altered gravity on the development of the
nance of cell architecture, cell motility, cell divi- neurovestibular system in aquatic vertebrates
sion, enzyme regulation, ion channels, and gene have been found, especially regarding the neural
expression. In micro-/hypergravity the gene control of inner ear otolith mineralization in fish
expression profile of pro- and eukaryotic cells (Fig. 1). A closer understanding of the reasons for
change. These alterations involve numerous (human) motion sickness susceptibility was
important cellular pathways. Much progress gained by these experiments. In general, the find-
has been made in the understanding of ings support the “otolith asymmetry hypothesis”
Gravitational Biology, Fig. 1 Otolith mineralization is at Earth gravity (left image, 1 g), but did not grow that
affected by gravity. Note that a fish otolith grown 3 weeks large under hypergravity (1 g, left: 175 mm; hg, right:
at 3 g hypergravity (right image, hg) has a core (indicating 115 mm) (Photo: M. Beier)
the begin of the experiment) as small as in a control reared
1006 Gravitational Biology
dependent displacement of statoliths (small

grains of barium sulfate), which get into contact
with gravireceptor proteins in the lower cell
membrane. Gravireceptor activation in
gravistimulated rhizoids induces a local reduc-
tion of cytosolic calcium at that area where the
statoliths are sedimented. This causes a local
inhibition of exocytosis which results in differen-
tial extension of the opposite cell flanks and the
gravitropic response of the rhizoid, namely, the
correction of the growth direction.
In recent years Arabidopsis thaliana has
become the model system of choice to examine
gravity-related responses and regulation in higher
plants due to the availability of genomic infor-
mation. While the gravitropism of roots and
shoots of higher plants is well known for over a
century a detailed understanding of the signal
perception and transduction has not yet been
Gravitational Biology, Fig. 2 How gravity shapes life achieved. Already Darwin identified the root
morphology (Photo: P. Richter) cap as well as starch containing particles
(amyloplasts which serve as statoliths in the root
cap) as important for gravitropism. Another line
which relates motion sickness to differential sen- of evidence showed that a phytohormone, auxin,
sory input from the left and the right inner ear. is involved, too. A differential displacement of
Thus, numerous experiments on fish as model auxin was identified as main cause for the
system have shown that particular neuroscientific reorientational growth of plant organs in respect
issues can be transferred to human physiology. to gravity (but also to other stimuli like light).
The current model of gravitropism is based on the
Plant Biology differential transport of auxin due to
Plants demonstrate in a fascinating and easily specific membrane proteins (PIN proteins).
detectable way the impact of gravity on biologi- In roots, auxin is transported in the direction of
cal systems (Fig. 2). Considerable progress on the root caps by the combined action of several
basic knowledge of plant gravity sensing and PIN proteins. The location of one of the PIN
gravitropism has been achieved regarding espe- proteins in the membrane changes according to
cially the basic knowledge on the effects of the orientation of the root. As a result the
altered gravity (gravitational stress) on essential concentration of the auxin increases in the lower
cellular functions related to plant growth and half of a gravistimulated root. However, this
development. Experiments performed in micro- model includes many unproven hypotheses
gravity have greatly contributed to the under- and the current view might well change in the
standing of how plants sense the direction of future.
gravity and respond to it. As a simple unicellular
system the green algae Chara shows very pro-
nounced gravitropism (growth in relation to the Applications
gravity vector) which is based on the directional
growth of the opposite flanks to the cell tip. The Application of gravitational biology aims at the
tip growth of the cell is controlled by the gravity solution of terrestrial and space-related problems,
with the latter occurring during space flight. Iden- ▶ Microgravity

tified issues include the progressive atrophy of ▶ Microorganism
bones and muscles as well as the lack of stimula- ▶ Organelle
tion of immune cells. The recent attempts to ▶ Planetary and Space Simulation Facilities
understand the microgravity-induced alterations ▶ Radiation Biology
on the molecular and cellular level increase our ▶ Space Biology
knowledge in basic science and might well con-
tribute to human health considerations, e.g.,
depression of the immune system, induction of References and Further Reading
cancer, muscle atrophy, osteoporosis which are
serious health issues on long-term space flights, Anken R (2006) On the role of the central nervous system
but which are also relevant for health problems in regulating the mineralisation of inner-ear otoliths of
on Earth. Understanding of the molecular and fish. Protoplasma 229:205–208
Braun M, Limbach C (2007) Rhizoids and protonemata of
cellular basis of gravitropic mechanisms in plants characean algae: unicellular model systems for
G
is considerably important for many aspects of research on polarized growth and plant gravity sens-
optimization of plant breeding and agriculture ing. Protoplasma 229:133–142
on Earth but also with respect to growing of Brinckmann E (ed) (2007) Biology in space and life on
Earth, effects of spaceflight on biological systems.
plants in space (Space Farming). Wiley-VCH, Weinberg
Clément G, Slenzka K (eds) (2006) Fundamentals of space
biology. Microcosm Press/Springer, New York
Future Directions Cogoli A (ed) (2002) Cell biology and biotechnology in
space. Elsevier, Amsterdam
Häder D-P, Hemmersbach R, Lebert M (2005) Gravity
In recent years research directions changed from and the behaviour of unicellular organisms. Cam-
pure descriptive assemblies of observations to a bridge University Press, Cambridge
molecular understanding of the underlying mech- Häder R, Richter P, Schuster M, Daiker V, Lebert
M (2009) Molecular analysis of the graviperception
anisms. New research directions are fostered by signal transduction the flagellate Euglena gracilis:
the availability of whole genomes and high- involvement of a transient receptor potential-like
throughput technologies (-omics). This includes channel and calmodulin. Adv Space Res
the long-term influence of microgravity on the 43:1179–1184
Horn E, Böser S, Membre H, Dournon C, Husson D,
development of animals and plants by means of Gualandris-Parisot L (2006) Morphometric investiga-
multigeneration experiments. These experiments tions of sensory vestibular structures in tadpoles
require biological life-support systems which (Xenopus laevis) after a spaceflight: implications for
allow the analysis of the complex interaction of microgravity induced alterations of the
vestibuloocular reflex. Protoplasma 229:193–203
different species in artificial habitats with the Horneck G, Baumstark-Khan C (eds) (2002) Astrobiology,
direct application in the future of food produc- the quest for the conditions of life. Springer,
tion, waste removal (solid, liquid, and gaseous), Heidelberg
and recycling. Knowing how organisms manage Morey-Holton E (2003) The impact of gravity on life. In:
Rothschild L, Lister A (eds) Evolution on planet
the absence of gravity is mandatory for long-term Earth – the impact of the physical environment. Aca-
orbital and interplanetary missions, but will well demic, New York, pp 143–159
have an impact on health and plant breeding Position Paper: European Science Foundation (2008) Sci-
issues on Earth. entific evaluation and future priorities of ESA’s ELIPS
programme. ESSC-ESF position paper 2008,
96 p. http://www.esf.org/research-areas/space/publi-
cations.html
See Also Teale WD, Ditengou FA, Dovzhenko AD, Li X,
Molendijk AM, Ruperti B, Paponov I, Palme
K (2008) Auxin as a model for the integration of
▶ Gravitation hormonal signal processing and transduction. Mol
▶ International Space Station Plant 1(2):229–237
1008 Gravitational Collapse, Planetary
Gravitational Collapse, Planetary ▶ Protoplanetary Disk Instability
▶ Star Formation, Theory
Sean N. Raymond ▶ Star Formation, Triggering
Boss A (1997) Giant planet formation by gravitational

Definition instability. Science 276:1836–1839
Johansen A et al (2007) Rapid planetesimal formation
Gravitational collapse is the collapse of a region of in turbulent circumstellar disks. Nature
448:1022–1025
material under its own gravity, for example, of the
dense core of an interstellar cloud on its way to
becoming a star. Gravitational collapse occurs
when the local self-gravity (i.e., the inward-
Gravitational Collapse, Stellar
directed force) exceeds the restoring force, which
is often in the form of thermal gas pressure or
Steven W. Stahler
turbulent stirring.
Overview
Definition
In astrophysics and planet formation, gravita-
tional collapse is extremely important, and the
The rapid contraction of a fluid mass because of
process is thought to occur in a variety of settings
the mutual gravitational attraction of its compo-
and on a range of different size scales. For exam-
nent particles is called gravitational collapse. If
ple, stars are thought to form from clumps of gas
the forces supporting the object are weak, then its
that are gravitationally unstable within ▶ molec-
internal elements freely fall toward one another,
ular clouds. In addition, an alternate to the core
leading to a highly condensed final configuration.
accretion model for giant planet formation is the
As a consequence of the inverse-square nature of
top-down, disk instability model. This model pro-
the gravitational force, the duration of collapse
poses that giant planets may form within the very
(known as the ▶ free-fall time) depends mainly
short span of a few thousand years in the outer
on the object’s initial density. In practice, only
regions of gravitationally unstable ▶ protoplane-
the very large masses found in astronomical bod-
tary disks. Finally, models for the formation of
ies are susceptible to this process. Examples are
▶ planetesimals suggest that they may form via
dense cores within molecular clouds, which col-
the gravitational collapse of aggregates of
lapse to form stars, and the interiors of massive
pebble- to boulder-sized bodies that are concen-
stars, prior to supernova explosion.
trated within turbulent structures in protoplane-
tary disks.
See Also
See Also ▶ Fragmentation of Interstellar Clouds

▶ Free-Fall Time
▶ Core Accretion, Model for Giant Planet ▶ Protostars
Formation ▶ Protostellar Envelope
▶ Planetesimals ▶ Star Formation, Theory
Great Oxygenation Event 1009
Gravitational Focusing Gravity
Rory Barnes ▶ Gravitation

Washington, Seattle, WA, USA
Great Oxidation Transition

Definition
▶ Great Oxygenation Event
Gravitational focusing refers to the enhancement
in the likelihood that two particles will collide,
due to their mutual gravitational attraction. With-
Great Oxygenation Event G
out gravity, the probability of a collision scales
with the cross-sectional area of the two particles,
Andrey Bekker
but with gravity, some particles that would have
Department of Geology and Geophysics, Yale
missed each other can still collide. In other
University, New Haven, CT, USA
words, the particles have effective cross sections
larger than their physical cross sections. In the
presence of only two bodies (two-body approxi-
Keywords
mation), the gravitational enhancement factor is
given by 1 + (vesc /vrel)2, where vesc is the parti-
Paleoproterozoic; Lomagundi carbon isotope
cle’s escape speed and vrel is the relative velocity
excursion; Huronian glaciation; Rise of atmo-
of the two particles. Gravitational focusing is
spheric oxygen; Great oxidation event; Great oxi-
important during planet formation, as it leads to
dation transition
faster growth.
Synonyms
See Also
Great oxidation transition; Oxyatmoversion, rise
of atmospheric oxygen
▶ Escape Velocity
Definition
Gravitational Instability The Great Oxidation Event (GOE) refers to the

transition from the mildly reducing Archean
▶ Protoplanetary Disk Instability atmosphere-ocean system to the oxygenated
atmosphere and shallow oceans of the early
Paleoproterozoic that started between ~2.4 and
2.3 Ga and ended between 2.1 and 2.0 Ga
(Holland 2002). The beginning and the end of
Gravitational Instability (Model for the GOE were never explicitly defined. In general
Giant Planet Formation) terms, the beginning of the GOE is marked by the
disappearance of mass-independent fractionation
▶ Disk Instability, Model for Giant Planet of S isotopes and an increased range of mass-
Formation dependent fractionation of sulfur isotopes in
1010 Great Oxygenation Event
association with highly variable carbon isotope strong support for atmospheric oxygenation in
record, Huronian glaciations, and the final stage the Middle Precambrian. S.M. Roscoe (1969,
in the assembly of the supercontinent (Bekker 1973) working for the Geological Survey of Can-
et al. 2004; Holland 2006; Bekker and Holland ada in the Huronian basin of Southern Ontario
2012). Although the term implies a relatively observed that the lower part of the Huronian
short duration, as that of phenomena used in succession contains detrital pyrite and uraninite
event stratigraphy, the GOE is originally defined and reduced paleosols, while the upper part has
to last more than 200 Ma and finish at the end of red beds and oxidized paleosols; sandwiched
the >2.22–2.06 Ga Lomagundi carbon isotope between these levels, the middle part recorded
excursion during which between ~12 and three glacial events (Fig. 1). Similar rounded
22 times the oxygen content of the present-day grains of pyrite, uraninite, and siderite (Fig. 2),
atmosphere was released to surface environments which are not stable in contact with oxidized
(Karhu and Holland 1996). While the internal waters and oxygenated atmosphere, were also
texture of the GOE is still unresolved and remains observed worldwide in fluvial conglomerates
to be a topic of further work, during these from sedimentary successions of similar and
200 Ma, atmospheric oxygen likely rose from older age (e.g., Schidlowski 1966; Rasmussen
the level of <105 PAL (Present Atmospheric and Buick 1999). Roscoe (1973) named this tran-
Level) to a magnitude of fully oxygenated atmo- sition at ~2.3 Ga “oxyatmoversion.” Interest-
sphere (likely below the modern level) before ingly, as shown on Fig. 1, more than 40 years
rapidly falling to much lower concentrations in later, our geochronological and stratigraphical
the immediate aftermath of the Lomagundi car- estimates for this transition did not change sig-
bon isotope excursion. The GOE ended up with nificantly. Preston E. Cloud of the University of
the ocean deoxygenation in association with the California, Los Angeles (UCLA), linked biolog-
supercontinent breakup, sea-level rise, and depo- ical evolution and environmental changes
sition of Mn- and P-rich sediments. suggesting that once efficient oxygen- and
peroxide-mediating enzymes developed, oceans
became oxygenated and banded iron formations
History were replaced by red beds in the rock record by
~1.8–2.0 Ga (Cloud 1968). Heinrich D. Holland
The notion that the early Earth’s atmosphere did of Princeton University, on the basis of geochem-
not contain oxygen is deeply rooted in studies of ical arguments and the presence of detrital urani-
biology and chemistry in the eighteenth and nine- nite in siliciclastic fluvial and shallow-marine
teenth centuries (Priestley 1781; Phipson 1893). successions, inferred the occurrence of a largely
The first detailed discussion, founded on geolog- anoxic atmosphere before ~1.8 Ga (Holland
ical evidence that atmospheric oxygen was low 1962). These interpretations were not universally
and then rose in the Middle Precambrian, was accepted and they were contested on various
presented by MacGregor (1927), a director of occasions (e.g., Dimroth and Kimberley 1976;
the Rhodesia Geological Survey, based on his Ohmoto 2004 and references therein). Arguments
observations on sedimentary successions pre- of the 1970s–1990s helped to deepen our under-
served in the Archean ▶ greenstone belts and standing of timing and different geochemical and
the Paleoproterozoic Magondi Mobile Belt of geological proxies for the rise of atmospheric
Rhodesia (Zimbabwe). This interpretation oxygen as well as to quantify atmospheric and
remained largely unnoticed since poor age con- seawater oxygen levels implied by these proxies.
straints limited correlation of Precambrian sedi- Most of these arguments were put to rest by 2000,
mentary successions, making it difficult to when James Farquhar of the University of Mary-
constrain the age and stratigraphic position of land (Farquhar et al. 2000) discovered mass-
this event. By the 1960s, several independent independent fractionation of sulfur isotopes in
lines of evidence were employed to provide a Archean sedimentary sulfides and sulfates,
Great Oxygenation Event, Fig. 1 Age-calibrated strat- et al. (2013) for references) and the age of the Copper Cliff
igraphic column of the Paleoproterozoic Huronian Super- rhyolite at the base of the Huronian Supergroup (Ketchum
group, Ontario, Canada (Updated from Rasmussen et al. 2013). Values in blue adjacent to marine carbonate
et al. (2013)). Dates for tuff beds of the Gordon Lake units represent their carbon isotope compositions. MDF is
Formation are from Rasmussen et al. (2013); also shown mass-dependent fractionation, MIF is mass-independent
is the age of the Nipissing dyke swarm (see Rasmussen fractionation of sulfur isotopes, respectively
which can be only explained by photochemical low-level oxygenation with a geologically rapid
processes in the atmosphere with low levels of transition to the oxygenated atmosphere at
oxygen (<105 PAL). This approach was subse- ~2.4 Ga.
quently applied to date the rise of atmospheric
oxygen to between ~2.45 and 2.32 Ga (Bekker
et al. 2004; Hannah et al. 2004; Guo et al. 2009). Overview
In the last decade, a number of studies were
published, which suggested transient, low-level Evidence is growing for supporting oxygenic
oxygenation events before the GOE, of either photosynthesis as early as ~3.0 Ga ago and for
global or local extents (e.g., Anbar et al. 2007; transiently, globally or locally, oxygenated envi-
Crowe et al. 2013; Planavsky et al. 2014). The ronments since that time (Anbar et al. 2007;
current and future research is now faced with a Crowe et al. 2013; Planavsky et al. 2014). How-
challenge: how to reconcile the notion of the ever, it is clear that mass-independent fraction-
long-lasting (>1 Ga) period with transient or ation of sulfur isotopes persisted in the
Great Oxygenation Event, Fig. 2 Mineralogical and rounded siderite grain, with core of compositionally
lithological proxies for Archean anoxic atmosphere and banded siderite and syntaxial overgrowth (From Rasmus-
ocean: (a) detrital, muffin-shaped grains of uraninite with sen and Buick 1999); (c) detrital, rounded pyrite grains
galena overgrowths from Archean fluvial conglomerates from the ~2.64 Ga Black Reef Quartzite, South Africa; (d)
of South Africa (From Schidlowski 1966); (b) detrital, ~2.45 Ga banded iron formation, Western Australia
sedimentary record until ~2.32 Ga, indicating et al. 2013). Association between the rise of
that oxygen was not a stable component of the atmospheric oxygen and global glaciations in
atmosphere at concentrations higher than 105 the Paleoproterozoic has been explained by high
PAL, until more than half billion years later (>1,000 ppm) concentrations of methane in the
(Farquhar et al. 2000; Bekker et al. 2004). This Archean anoxic atmosphere, contributing to
is somewhat disconcerting considering a rela- greenhouse conditions (Pavlov et al. 2000). The
tively small size of atmospheric oxygen reservoir rise of atmospheric oxygen would decrease meth-
and theoretical modeling indicating bistability of ane residence time and its concentration in the
oxygen in the atmosphere at either very low atmosphere, resulting in loss of greenhouse con-
(<105 PAL) or substantial (>5 103 PAL) ditions; the Huronian ice ages in this scenario
steady-state concentrations (Goldblatt could reflect a transition from the anoxic,
et al. 2006). The rise of atmospheric oxygen in methane-rich Archean atmosphere to the oxygen-
the early Paleoproterozoic is associated with the ated atmosphere with much lower levels of meth-
Huronian ice ages, when the Earth experienced ane and sufficient concentrations of CO2 to
four, potentially low-latitude glaciations between maintain greenhouse conditions (Bekker and
~2.45 and 2.22 Ga ago (Fig. 3; Rasmussen Kaufman 2007).
Mn-rich formations
Iron formations
Paleosols
Red beds
Detrital pyrite and uraninite
Marine sulfate evaporites
δ34 S record
Δ33 S record
1 2 3 4 Lomagundi Glaciations
10 Event (LE)
13C(‰,V-PDB)
0 G
Great Oxidation
Event (GOE)
−5
2.5 2.4 2.3 2.2 2.1 2.0
Age (Ga)
Great Oxygenation Event, Fig. 3 Secular carbon iso- after ca. 2.07 Ga indicates that sulfate evaporites again
tope variations in seawater and redox indicators for the became rare in the Paleoproterozoic and Mesoproterozoic
oxidation state of the early Paleoproterozoic atmosphere- records after that time. Deposition of iron formations and
ocean system (Modified from Bekker and Holland 2012). Mn-rich deposits indicates anoxic conditions in deepwa-
Four blue vertical bars mark Paleoproterozoic glacial ters. While deposition of iron formations does not neces-
events; the dashed secular carbon isotope curve between sarily require atmospheric oxygen and can be mediated by
2.5 and 2.22 Ga emphasizes the uncertainty in this part of anoxygenic photosynthetic bacteria, Mn oxidation
the curve; the dashed bar for marine sulfate evaporites requires significant levels of atmospheric oxygen
The rise of atmospheric oxygen is also associ- metals such as U, Mo, and Re in organic-matter-
ated with dramatic fluctuations, with escalating rich shales appear at that time, indicating expan-
magnitudes, in biogeochemical carbon cycle fol- sion of oxidized environments. The end of the
lowing the long-term stability of the Archean Lomagundi carbon isotope excursion is brack-
carbon cycle (Fig. 3; for notation, see ▶ Delta, eted between 2.11 and 2.06 Ga (Karhu and Hol-
Isotopic); their development immediately before land 1996) and marked by deposition of organic-
and during the Huronian ice ages argues for a matter-rich shales with highly 13C-depleted iso-
dynamic state of carbon cycle. The carbon isotope values and Mn- and P-rich sediments,
tope values of carbonates are mostly at around +8 whereas sulfate evaporites became again scarce,
‰ for the duration of the >2.22–2.06 Ga and proxies for ocean oxidation state indicate
Lomagundi carbon isotope excursion (Bekker expansion of anoxia at the end of the GOE (e.g.,
et al. 2003; Fig. 3). Since positive C-isotope Scott et al. 2014).
values in carbonates are thought to reflect high The beginning of the Lomagundi carbon iso-
relative burial rate of organic carbon, between tope excursion has been related to high continen-
12 and 22 times, the present atmospheric oxygen tal nutrient (e.g., phosphorus) flux linked to
inventory could have been released to the surface acidic groundwaters during the GOE
environments during the Lomagundi carbon iso- (Konhauser et al. 2011; Bekker and Holland
tope excursion (Karhu and Holland 1996), with 2012). The high acidity in terrestrial waters was
iron and sulfur in the continental crust consuming related to extensive oxidation of sulfides for the
most of it. Indeed, red beds, sulfate evaporites, first time in Earth’s history during continental
and high concentrations of redox-sensitive trace weathering. Acidic conditions in continental
runoff promoted dissolution of phosphates and oxygen sinks in the surface environments or
higher P delivery to the ocean resulting in higher increase oxidation potential of volcanic gases
relative burial rate of organic carbon and associ- generated via crustal subduction and recycling
ated oxygen release. The duration of the (Holland 2009; Claire et al. 2006; Catling
Lomagundi carbon isotope excursion is compa- et al. 2001; Kasting 2013; Zahnle et al. 2013).
rable to the half-life of sedimentary rocks in the All these models are concerned with the changes
continental crust suggesting that groundwater in the size of oxygen sinks and assume that the
acidity could have been largely titrated by the oxygen source did not change in size since oxy-
end of the excursion. This, by itself, could have genic photosynthesis evolved.
resulted in decreased nutrient flux to the ocean An alternative could be that oxygenic photo-
ending the GOE (cf., Bekker and Holland 2012). synthesis was limited by low continental flux of
Alternatively, weathering of organic-matter-rich nutrients (e.g., Mo and V; important for nitrogen
shales deposited during the Lomagundi carbon fixation) into the Archean ocean, resulting in
isotope excursion and uplifted during the limited net primary productivity and organic car-
~2.1 Ga orogenic events could have contributed bon burial. This suggestion is generally dismissed
reduced carbon to the atmosphere-ocean system since it would have significant repercussions for
and resulted in expansion of anoxic settings (cf., the biogeochemical carbon cycle, which are not
Kump et al. 2011; Bekker and Holland 2012). apparent in the carbon isotope record of Archean
Although it remains unclear which of these two carbonates (e.g., Kasting 2013). However, conti-
scenarios (or both) ended the GOE, the largest nental weathering of sedimentary rocks would
and longest-lived carbon isotope excursion in not return buried organic carbon to biogeochem-
Earth’s history ended up with the return to ical cycling under the Archean anoxic atmo-
low-oxygen levels in the atmosphere-ocean sphere; rather it would be buried again as
system. recycled organic matter with sediments. Simi-
While the timing and texture of the GOE is larly, carbon isotope mass balance requires that
progressively better understood, mechanisms for carbon released by dissolution of sedimentary
delayed oxygenation remain highly speculative. carbonates was also excluded from biogeochem-
The proposed diverse mechanisms can be broadly ical carbon cycle to satisfy carbon isotope values
grouped into three categories relying on of Archean carbonates near to 0 ‰. Conse-
(1) changes in mantle properties, (2) continental quently, carbon isotope records are actually con-
crustal growth, and (3) progressive oxidation of sistent with limited net primary productivity and
continental or oceanic crust. Inferred changes in organic-carbon burial under the Archean anoxic
mantle properties include oxidation of the mantle atmosphere-ocean conditions (Fig. 4).
by crustal subduction and mantle overturn (Kump The critical gap in our understanding of the
et al. 2000; Holland 2002) and progressive GOE is why it occurred between ~2.45–2.32 Ga.
cooling of the upper mantle (Konhauser Association with the supercontinent assembly
et al. 2009). Continental crustal growth is hold and 2.45–2.5 Ga mantle plumes has been previ-
responsible for sea-level fall and increased sub- ously recognized (e.g., Barley et al. 2005; Kump
aerial volcanism during the Archean to Paleopro- and Barley 2007), but was not satisfactorily
terozoic transition, which resulted in delivery of explained. In view of the above discussed limited
more oxidized volcanic gases with respect to net primary productivity and nutrient limitation
predominantly submarine volcanism in the in the Archean oceans, it is worth considering
Archean (Kump and Barley 2007; Gaillard enhanced continental nutrient flux under highly
et al. 2011). Progressive oxidation of oceanic or acidic groundwater conditions during mantle
continental crust by either hydrogen escape from plume events, when large amounts of CO2 and,
the atmosphere or oxygenic photosynthesis is especially, SO2 were delivered to the anoxic
inferred in a number of models to either decrease atmosphere and precipitated as acidic aerosols
the relationships among the demise of MIF in

S isotopes in the sedimentary record, evidence
for atmospheric and ocean oxygenation, ice
ages, and dynamic state of biogeochemical car-
bon cycle clearly argue against this delay (Fig. 3).
As our understanding of Earth’s history evolves
and as we answer some of our questions, more
complex puzzles surface and motivate us to
search for a deeper grasp. We are reasonably
confident now when the GOE happened and
what were other events associated with it. The
questions that we will be likely faced for another
decade or two are these: Why was the GOE
delayed? Why did it happen when it happened? G
Why didn’t biogeochemical carbon cycle change
after the GOE? To answer these questions, a
combination of theoretical modeling with field
Great Oxygenation Event, Fig. 4 Archean biogeo-
chemical carbon cycle; fluxes for the Neoarchean are in and laboratory studies will be required. The pro-
1012 mol/year from Bekker and Holland (2012). Isotopic spective of full understanding of the Earth system
composition of Archean carbonates and organic matter is is as ever a tantalizing goal to reach.
based on the geological records. It is proposed that organic
and carbonate carbon contents of the Archean crust have
been growing, resulting in a progressively increasing sub-
duction flux of carbon to the mantle. Both carbonate and
organic carbon were recycled during continental See Also
weathering, but did not pass through biogeochemical car-
bon cycle due to the lack of oxygen in the Archean atmo-
sphere and low flux of nutrients to the oceans (see text for ▶ Cap Carbonates
more details) ▶ Carbon Cycle, Biological
on the continents. Similar scenario could poten- ▶ Diamictite/Diamicton
tially explain Archean transient oxidation events, ▶ Fractionation, Mass Independent and
but in contrast to these short-lived events, the Dependent
2.45–2.5 Ga time interval is marked by multiple ▶ Glaciation
mantle plume events that affected all continents. ▶ Isotope Biosignatures
This period was previously singled out in the ▶ Oxygenation of the Earth’s atmosphere
Earth’s history and was linked to a fundamental ▶ Paleosols
change in heat flux at the core-mantle boundary ▶ Precambrian
(Heaman 1997). ▶ Proterozoic Eon
The recent model for recycling mass-indepen- ▶ Pyrite
dent fractionation of sulfur (MIF-sulfur) from the ▶ Red Beds
continental crust to the oceans allows for as much ▶ Sedimentary Rock
as 100 Ma delay for the disappearance of MIF ▶ Snowball Earth
from the sedimentary record after the GOE ▶ Stable Isotopes
started (Reinhard et al. 2013). In this model, ▶ Stratigraphy
atmosphere could have had significant levels of ▶ Sulfur
oxygen resulting in continental weathering under ▶ Sulfur Cycle
oxic conditions, while seawater sulfate and sedi- ▶ Transvaal Supergroup, South Africa
mentary sulfur would still carry MIF. However, ▶ Weathering
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Anbar AD, Duan Y, Lyons TW, Arnold GL, Kendall B, ogy 25:299–302
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Garvin J, Buick R (2007) A whiff of oxygen before atmosphere. In: Engel AEJ, James HL, Leonard BF
the great oxidation event? Nature 317:1903–1906 (eds) Petrologic studies – a volume to honor
Barley ME, Bekker A, Krapez B (2005) Late archean to A. F. Buddington. Geological Society of America,
early paleoproterozoic global tectonics, environmental Denver, pp 447–477
change and the rise of atmospheric oxygen. Earth Holland HD (2002) Volcanic gases, black smokers, and
Planet Sci Lett 238:156–171 the great oxidation event. Geochim Cosmochim Acta
Bekker A, Holland HD (2012) Oxygen overshoot and 66(21):3811–3826
recovery during the early Paleoproterozoic. Earth Holland HD (2006) The oxygenation of the atmosphere
Planet Sci Lett 317–318:295–304 and oceans. Phil Trans R Soc A 361:903–915
Bekker A, Kaufman AJ (2007) Oxidative forcing of global Holland HD (2009) Why the atmosphere became oxygen-
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Green Bank Equation 1017
Planavsky NJ, Asael D, Hofmann A, Reinhard CT, Definition

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See Also
Green Bacteria
▶ Carbon Dioxide
▶ Citric Acid Cycle
Ricardo Amils
▶ Photoautotroph
Synonyms Green Bank Equation
Green nonsulfur bacteria; Green sulfur bacteria ▶ Drake Equation

1018 Green Nonsulfur Bacteria
loss of hydrogen to space (e.g., a possible sce-

Green Nonsulfur Bacteria nario for Venus, if it had a similar water reservoir
as the Earth).
▶ Green Bacteria
See Also
Green Rust ▶ Absorption Cross Section

▶ Iron Oxides, Hydroxides and Oxy-hydroxides ▶ Grey Gas Model
▶ Habitable Zone
▶ Non-grey Gas Model: Real Gas Atmospheres
Green Sulfur Bacteria
▶ Green Bacteria
Greenland
Minik T. Rosing
Greenhouse Effect
Nordic Center for Earth’s Evolution, Natural
History Museum of Denmark, University of
Lisa Kaltenegger
Copenhagen, Copenhagen, Denmark
Definition
Definition
Greenland is a large island in the North Atlantic
The warming of a planet by the “greenhouse
between Europe and North America, part of the
effect” is caused by atmospheric gases that are
Nain craton. It is composed dominantly of Precam-
very efficient absorbers in the infrared but not in
brian rocks, but these are exposed and accessible
the visible. Symmetric, nonpolar molecules such
only along the coastlines at the margins of the central
as H2, N2, and O2 have no electric dipole and
ice cap. The early Archean terranes near the town of
cannot absorb or emit photons. They are therefore
Nuuk on the SW coast include some of the oldest
not greenhouse gases. The visible light of the
rocks on Earth. In the Isua belt are found ca. 3.8 Ga
parent star reaches and heats the planetary sur-
old metamorphosed pillow basalts and clastic and
face and is reemitted in the infrared, where part of
chemical sediments that constitute the oldest known
the energy is absorbed by the atmospheric green-
supracrustal sequences. Fractionated C, N, and
house gases with some radiated back toward the
S isotopic compositions of materials of probable
surface. Water, carbon dioxide, and methane are
sedimentary origin are thought to be biogenic and
the greenhouse gases that raise the surface tem-
to provide the oldest record of life on Earth.
perature on Earth to an average of +15 C instead
of 17 C. The “runaway greenhouse” occurs
when greenhouse conditions vaporize part of a
water reservoir, the resulting water vapor See Also
increases the greenhouse effect, and this feed-
back cycle leads to loss of the entire surface ▶ Akilia
water and, as a secondary effect, to the photodis- ▶ Earth, Formation and Early Evolution
sociation of the atmospheric water vapor and the ▶ Isua Supracrustal Belt
Greenstone Belts 1019
granite and gneiss, are the constituents of

Greenschist Facies Archean and Proterozoic cratons.
The volcanic rocks range in composition from
Nicholas Arndt ▶ komatiite to rhyolite but are dominated by
ISTerre, Université Grenoble Alpes, France ▶ basalt; sedimentary rock types are diverse and
include shale, siltstone, sandstone, conglomerate,
rare carbonates and evaporites, and (bio)chemical
Definition precipitates such as banded-iron formation and chert.
The geochemical composition of the basaltic
Greenschist facies is one of the major subdivisions rocks shows that some formed as ▶ oceanic crust,
within the classification scheme of metamorphic perhaps as part of oceanic plateaux, and that
rocks. Greenschist facies refers to low- to others formed in island arcs. The sedimentary
medium-grade metamorphic conditions rocks usually document the evolution of green-
corresponding to temperatures of about stone belts from deep-marine, open-oceanic set- G
300–500 C and pressures of 3–20 kbar, tings through marginal basins filled with
corresponding to crustal depths of about 8–50 km. voluminous turbidites and chemical sediments.
The name is derived from the green color of its Very few greenstone belts include a sedimentary
characteristic minerals (chlorite) and the schistose cap of shallow-water and quartz-rich sediments.
fabric induced by recrystallization and deformation. Although the metamorphic grade of most
Characteristic minerals are chlorite, actinolite, and greenstone belts ranges from sub-greenschist to
albite epidote in metabasalt; chlorite, serpentine amphibolite facies, ▶ greenschist-facies metamor-
talc, tremolite, and brucite in meta-ultramafic phism dominates. Metamorphic grade typically
rocks; and muscovite, garnet, and andalusite in increases toward the plutonic or gneiss terranes
metapelitic sediments. Greenschist facies prevails bordering the greenstone belts and toward internal
in the middle levels of oceanic crust and in the upper ductile shear zones. Abundant chlorite and actin-
levels of orogenic belts. Many supracrustal rocks in olite in ▶ greenschist-facies basalts and
Archean terranes are metamorphosed at greenschist serpentinite formed from ultramafic rocks impart
facies, hence the name “▶ greenstone belts.” a green color that gives rise to the name.
Because greenstone belts can be considered the
down-sagged “keels” of deformed sedimentary
See Also
basins, most units in greenstone belts dip steeply,
are tightly folded, and are cut in places by large shear
zones. These developed as parts or all of the volcano-
▶ Metamorphism
sedimentary sequence were accreted to a continent
or deformed in a continental collision zone.
Archean greenstone belts represent the oldest
record of Earth’s surface conditions, including
Greenstone Belts the evolution and spreading of ▶ life on Earth.
They host important ore deposits of metals such
Nicholas Arndt as gold, Cu-Zn, Ni, and Fe. Well-known exam-
ISTerre, Université Grenoble Alpes, France ples include the Abitibi and Flin Flon belts in
Canada, the Norseman-Wiluna belt in Australia,
and the Barberton, Sutherland, and Murchison
Definition belts in South Africa.
A greenstone belt is a usually elongate structure See Also

composed dominantly of metamorphosed volca-
nic and ▶ sedimentary rocks that, together with ▶ Barberton Greenstone Belt
1020 Grey Body
▶ Canadian Precambrian Shield complicates the solution of the radiative transfer

▶ Greenschist Facies equations, since the fluxes must be sought indi-
▶ Metamorphic Rock vidually on a very dense grid of wavelength,
▶ Metasediments and then the results integrated to yield the net
▶ Pilbara Craton atmospheric heating. The radiative transfer
equations become much simpler if the optical
thickness is independent of wavelength, which
is known as the grey gas approximation. For
grey gases, the relevant equations can be inte-
Grey Body
grated over wavelength, yielding a single differ-
ential equation for the net upward and downward
Daniel Rouan
flux. With the exception of clouds of strongly
absorbing condensed substances like water, the
Meudon, France
grey gas model yields a poor representation to
radiative transfer in real atmospheres, but the
grey gas model is nonetheless a powerful theo-
Definition
retical tool for exploratory work in planetary
atmospheres. It is also valuable as a radiation
A grey body is a hypothetical source of electro-
scheme in idealized general circulation model-
magnetic radiation that would emit as a ▶ black
ling studies.
body but with an ▶ emissivity lower than 1 and
constant with wavelength. The grey body
assumption is often more useful as a first approx-
imation to the actual emission of a real object or
See Also
medium than is the ideal black body radiation. It
is used in ▶ radiative transfer problems, for
instance, in planetary atmospheres.
▶ Exoplanets, Modeling Giant Planets’
See Also Atmospheres
▶ GCM
▶ Blackbody ▶ Habitable Planet, Characterization
▶ Emissivity ▶ Non-Grey Gas Model: Real Gas Atmospheres
▶ Radiative Processes ▶ Optical Depth
▶ Radiative Transfer

Grey Gas Model
Pierrehumbert RT (2010) Principles of planetary climate.
Ray Pierrehumbert Cambridge University Press, Cambridge
Department of the Geophysical Sciences,
University of Chicago, Chicago, IL, USA
Groove
Definition
▶ Rille
For most atmospheric gases the optical thickness ▶ Rima, Rimae
has an intricate dependence on wavelength. This ▶ Sulcus, Sulci
Gullies 1021
Definition
Guanine
On Earth, the term gully refers to a small and
Shin Miyakawa narrow but relatively deeply incised stream
Ribomic Inc., Minato-ku, Tokyo, Japan course. On ▶ Mars, the term is used somewhat
differently to denote a landform consisting
of – from top to bottom – (1) a catchment area
Definition or erosional alcove, (2) one or several transport
channels, and (3) a depositional fan or apron.
C5H5N5O. MW: 151.13. Guanine is one of four
nucleic acid bases found in DNA and ▶ RNA. In
double-stranded DNA, guanine base pairs with Overview
cytosine. It is hydrolyzed to xanthine. The half-
life to hydrolysis in solution is 0.8 years at 100 C Gullies were first unambiguously identified on G
and 1.3 106 years at 0 C at pH 7. It has a UV Mars in images taken by the Mars Orbiter Camera
absorption maximum at 246 and 276 nm (pH 7). (MOC) (Malin and Edgett 2000). Gullies are
It has been found in the Murchison meteorite and relatively small landforms, with common lengths
can be synthesized in HCN polymerizations, of a few kilometers only. Close morphological
Fischer-Tropsch type reactions, and electric dis- analogues to Martian gullies exist in the Arctic
charges acting on gas mixtures such as CO-N2- regions on Earth (e.g., Greenland, Svalbard), but
H2O. also in hot arid environments. On Earth, gullies
typically form by the action of liquid ▶ water.
The gullies on Mars are very young and could
have formed in the past several million years or
See Also
even more recently (Reiss et al. 2004). Most of
the gullies have been found in midlatitudes,
▶ Cytosine
where liquid surface water is physically not stable
▶ DNA
with the current Martian climate. Most of the
▶ Fischer-Tropsch-Type Reaction
present-day activity in gullies appears to be by
▶ HCN Polymer
mass wasting associated with CO2 frost/ice (Dun-
das et al. 2012). Many hypotheses have been put
▶ Murchison
forward to solve this conundrum, including a
formation by groundwater (Malin and Edgett
▶ RNA
2000), liquid CO2 (Musselwhite et al. 2001), or
dry granular flows (Treiman 2003). While these
models avoid the problem of forming midlatitude
liquid surface water in the recent past, they have
Gullies been challenged on the basis of physical and
morphological arguments (e.g., see the discus-
Ernst Hauber sion in Dickson and Head 2009). On the other
Deutsches Zentrum f€ur Luft- und Raumfahrt hand, scenarios involving liquid water as a result
(DLR) e.V., Institut f€ur Planetenforschung, of near-surface ground ice or snow melting
Berlin, Germany (Costard et al. 2002; Christensen 2003) would
imply precipitation (most probably snow) and,
therefore, a recent climate change on Mars. The
Keywords climate of Mars is susceptible to astronomical
forcing (i.e., the effect on climate of changes in
Erosion; Flooding; Groundwater; Ice; Water the tilt of the rotational axis of a ▶ planet and of
1022 Gunflint Formation
orbital parameters such as ▶ eccentricity) Musselwhite DS, Swindle TD, Lunine JI (2001) Liquid
(Jakosky et al. 1995), and it is plausible that CO2 breakout and the formation of recent small gullies
on Mars. Geophys Res Lett 28:1283–1286
periodic variations of the obliquity of the planet’s Reiss D, van Gasselt S, Neukum G, Jaumann R (2004)
spin axis in the past 105–106 years (Laskar Absolute dune ages and implications for the time of
et al. 2004) allowed the deposition of snow and formation of gullies in Nirgal Vallis, Mars. J Geophys
ice in midlatitudes (Mischna et al. 2003). The Res 109, E06007. doi:10.1029/2004JE002251
Treiman AH (2003) Geologic settings of Martian gullies:
possible existence of liquid water or water ice at implications for their origins. J Geophys Res
or near the surface classifies some gullies as 108:8031. doi:10.1029/2002JE001900
potential special regions that are banned from
being investigated in situ due to ▶ planetary pro-
tection rules.
Gunflint Formation
Takeshi Kakegawa1 and Akizumi Ishida2

See Also 1
Graduate School of Science, Tohoku University,
Sendai, Japan
▶ Eccentricity 2
Atmosphere and Ocean Research Institute,
▶ Mars
University of Tokyo, Kashiwa, Chiba, Japan
▶ Obliquity and Obliquity Variations
▶ Planet
Keywords
▶ Water
Gunflint Formation; Microfossils; Rove forma-
References and Further Reading tion; Stromatolites; Sudbury impact; Traces of
life
Christensen PR (2003) Formation of recent Martian
gullies through melting of extensive water-rich snow
deposits. Nature 422:45–48
Costard F, Forget F, Mangold N, Peulvast J-P (2002) For- Definition
mation of recent Martian debris flows by melting of
near-surface ground ice at high obliquity. Science The Gunflint Formation of the late Paleopro-
295:110–111 terozoic Animiki Group comprises chemical, clas-
Dickson JL, Head JW (2009) The formation and evolution
of youthful gullies on Mars: gullies as the late- tic, and volcaniclastic sedimentary rocks that are
stage phase of Mars’ most recent ice age. Icarus 100–160-m thick and which extend for 175 km
204:63–86 along the Gunflint Range from northeastern Min-
Dundas CM, Dieniega S, Hansen CJ, Byrne S, McEwen nesota to the Thunder Bay area of Ontario, Can-
AS (2012) Seasonal activity and morphological
changes in martian gullies. Icarus 220:124–143 ada. The Gunflint Formation is known for
Jakosky BM, Henderson BG, Mellon MT (1995) Chaotic occurrence of various ▶ microfossils, including
obliquity and the nature of the Martian climate. Gunflintia, Huroniospora, and Eoastrion. Fossil-
J Geophys Res 100:1579–1584 ized primitive fauna in Gunflint Formation is one
Laskar J, Correia ACM, Gastineau M, Joutel F, Levrard B,
Robutel P (2004) Long term evolution and chaotic of the best preserved in the geological record. For
diffusion of the insolation quantities of Mars. Icarus this reason, biogenicity of older microfossils and
170:343–364 dubiofossils is often tested by comparison with the
Malin MC, Edgett KS (2000) Evidence for recent ground- Gunflint Formation fossil record.
water seepage and surface runoff on Mars. Science
288:2330–2335
Mischna MA, Richardson MI, Wilson RJ, McCleese DJ Overview
(2003) On the orbital forcing of Martian water and
CO2 cycles: a general circulation model study with
simplified volatile schemes. J Geophys Res The Gunflint Formation was deposited on an
108:16–1. doi:10.1029/2003JE002051 Archean basement at the southwestern margin
Gunflint Formation 1023
of the Superior Province (see ▶ Canadian Pre- by an enhanced hydrothermal Fe flux, leading to
cambrian Shield). Occurrence of granular Fe-rich deep-water upwelling onto oxidized
peloidal and oolitic iron formations or taconite shelves. Previous studies also indicate the pres-
(an iron-bearing, high-silica, chert-like rock) ence of stratified ocean to form BIFs during depo-
characterizes the Gunflint Formation. Previous sition of the Gunflint Formation. Large quantities
investigators subdivided the Gunflint Formation of carbon isotope compositions of organic matter
into two members based on their respective and kerogen are available for the Gunflint and
lithologies (Fralick and Barrett 1995). The Rove Formations (Strauss and Moore 1992).
lower member typically has a basal conglomerate Most organic matter has d13C(PDB) values of
with algal chert, covered by cherty calcareous 31 to 36 ‰ (Fig. 1). Such values are consis-
rocks, changing into jaspilitic grainstone facies tent with carbon fixation via the Calvin cycle.
(see ▶ Jaspilite). The upper member has a stra- They are therefore consistent with earlier studies
tigraphy resembling that of the lower member: that assigned some of the Gunflint microfossils to
algal chert is followed by a succession of cherty the ▶ cyanobacteria phyla based on their mor- G
calcareous rocks and banded chert carbonate that phology (Barghoorn and Tyler 1965). Therefore,
passes westward into jaspilitic facies. Those it appears certain that oxygenic environments
facies suggest that the sediments of the Gunflint were widespread in the surface of the Gunflint
Formation were deposited under wave- Ocean as supported by the occurrence of banded
dominated and tide-dominated inter-shelf envi- iron formations.
ronments (Fralick and Barrett 1995). The carbo- Siliceous stromatolites in the Gunflint Forma-
naceous shales of the Rove Formation overlie the tion contain abundant microfossils and hematite.
upper member of the Gunflint Formation. Occur- Current Fe-isotope studies of hematite suggest
rence of various microfossils and ▶ stromatolites that such siliceous stromatolites formed in low
together with kerogenous sediments makes the oxygen conditions or near the redoxcline. This
Gunflint and Rove Formations attractive for engenders the possibility that a redoxcline was
astrobiologists to constrain the course of the present at shallow water depth, implying that
Paleoproterozoic ecological evolution reducing environments were more widespread
(Barghoorn and Tyler 1965). Accretionary lapilli among surface areas than was previously thought.
(rounded tephra balls consisting of volcanic ash The Fe-isotope studies suggest that chemoauto-
particles) were reported from near the Gunflint- trophic iron-oxidizing microbial ecosystems are
Rove boundary. These accretionary lapilli are potential primary producers at around the
often interpreted as impact-related ▶ ejecta redoxcline. Such geochemical data are consistent
(Addison et al. 2005). Such occurrence of accre- with some paleontological observations (Knoll
tionary lapilli creates another research interest for 2003).
astrobiologists, related to whether or not a Disagreement persists among researchers
Paleoproterozoic meteoritic impact might have (1) whether cyanobacteria were the major pri-
affected the microbial ecosystem. mary producers and (2) how deep the oxygenated
waters developed. Chemoautotrophic
Fe-oxidizing bacteria might be the alternative
Key Research Findings primary producers to cyanobacteria (Planavsky
et al. 2009). On the other hand, the activity of
A Redox-Sensitive Ecosystem chemoautotrophic Fe-oxidizing bacteria is
Banded iron formations (BIFs) disappeared after dependent on cyanobacterial activity because of
ca. 1.7 Ga. The BIFs of the Gunflint Formation their necessity of dissolved O2, which further
are thus among the last BIFs on Earth. It is gen- suggests that biomass production of chemoauto-
erally accepted that hydrothermal vent waters trophic Fe-oxidizing bacteria cannot exceed that
were the dominant source of dissolved iron to of cyanobacteria. Black shales of the Rove For-
form BIFs. The deep ocean becomes ferruginous mation are rich in organic carbon (up to 6 wt%),
1024 Gunflint Formation
Gunflint Formation,
Fig. 1 Carbon isotope
compositions of organic
matter and kerogen in
Gunflint and Rove
Formation
suggesting high microbial productivity during were limited in space. Therefore, the possibility
sedimentation. The absence of hematite deposi- exists that 13C-depleted organic matter and heavy
tion implies less influence of chemoautotrophic Fe-isotope in hematite might represent local
Fe-oxidizing bacteria in the Rove Formation than environmental effects rather than widespread
in Gunflint. In this situation, cyanobacteria are characteristics in the surface ocean.
the only reliable primary producers. ▶ Carbon
isotope compositions of organic matter of the Sudbury Impact Record
Rove Formation are compared to those of the The Sudbury Structure in Ontario (Canada) was
Gunflint Formation (Fig. 1). It is noteworthy most likely generated by a hypervelocity impact
that no carbon isotope difference was found at 1,850 1 Ma. Previous investigators searched
between organic matter of the Rove and Gunflint for “evidence” of the Sudbury impact in the Gun-
Formations, which suggests that the dominant flint Formation. The potential impact-related bed
primary producers for both formations were appears in the recrystallized and silicified carbon-
most likely cyanobacteria. ate sequence in the uppermost Gunflint Forma-
Microprobe analyses of iron were also tion. The proposed ejecta layer ranges from a few
performed on individual microfossils (House tens of centimeters thick (Fig. 2). Accretionary
et al. 2000). The Gunflint microfossils analyzed lapilli appear near the middle of this specific
using an ion microprobe show a range of layer. They are considered as ejecta products.
d13C(PDB) values of 32 to 45 ‰. Extremely Planar deformation features (PDFs) were also
13
C-depleted compositions down to 45 ‰ are found in quartz and feldspar grains. Most grains
not consistent with carbon fixation via the Calvin with PDFs are located within accretionary lapilli.
cycle, and they were explained by involvement of Tsunami-like deposits, associated with tektite
methanogens and methanotrophs in the ecosys- and glassy materials, also appear at the top of
tem. On the other hand, such 13C-depleted values the Gunflint Formation, for which the deposi-
were found only in microscale implying that tional age is consistent with the time of the Sud-
activities of methanogens and methanotrophs bury impact (Addison et al. 2005). Those features
Gunflint Formation 1025
Gunflint Formation,
Fig. 2 Accretionary lapilli
in Gunflint Formation
suggest that the Sudbury impact event was most to impact events. Furthermore, current experi-
likely recorded in the Gunflint Formation and that ments and theoretical studies suggest that impact
accretionary lapilli might represent impact ejecta, events can convert atmospheric CO2 and N2 into
although the alternative origins, such as volcanic CH4 and NH3, strongly affecting ecosystems and
ashes, are not excluded. Earlier reports described modifying the C and N isotope compositions in
the difference between the fossil-rich Gunflint the geological record. Therefore, C and N isotope
Formation and the fossil-absent Rove Formation compositions of ancient kerogens might be used
and discussed the possibility of impact-induced to evaluate the effect of an impact event on con-
extinction (Addison et al. 2005). temporary microbial ecosystem. In brief, no
marked change was observed in carbon isotope
compositions of organic matter from the Gunflint
Future Directions Formation to the Rove Formation (Fig. 1), which
implies that either (1) the Sudbury impact event
The late Paleoproterozoic represents a unique did not considerably affect the microbial commu-
period in the evolution of the redox conditions nities or (2) the impact event affected it, but the
of the atmosphere and oceans because very local- microbial ecosystem recovered quickly enough
ized oxic and anoxic conditions were present. that the influence was not recorded in rocks. To
During the transition from Gunflint to Rove evaluate the effect in greater detail, high-
oceans, the deep-water conditions became Fe resolution geochemical data will be necessary.
poor because of either euxinic or suboxic deep-
water conditions (Poulton et al. 2004; Slack
et al. 2007). Dynamics of redox changes and/or See Also
Fe-flux changes are important for the astrobiol-
ogy community to evaluate the tolerance and/or ▶ Banded Iron Formation
adjustability of anaerobic microorganisms to oxic ▶ Biomarker, Isotopic
and metal-poor conditions. The Gunflint and ▶ Carbon Isotopes as a Geochemical Tracer
Rove Formations will provide good opportunities ▶ Cyanobacteria
to examine the survival of microbial ecosystems ▶ Ejecta
1026 Gunflint Microbiota
▶ Impactite Definition
▶ Iron Isotopes
▶ Microfossils The Late Proterozoic ▶ Gunflint Formation com-
▶ Stromatolites prises 100–160 m thick iron-rich chemical, clas-
tic, and volcaniclastic sedimentary rocks
extending for 175 km along the Gunflint Range
from northeastern Minnesota, USA, to the Thun-
Addison WD, Brumpton GR, Valini DA, McNaughton der Bay area of Ontario, Canada. The iron forma-
NJ, Davis DW, Kissin SA, Fralick PW, Hammond tion is well dated at 1,878 1.3 Ma by U-Pb
AL (2005) Discovery of distal ejecta from the on zircons (Fralick et al. 2002). The microfossil
1850 Ma Sudbury impact event. Geology 33:193–196 assemblage of the Gunflint Formation is renowned
Barghoorn ES, Tyler SA (1965) Microorganisms from the
Gunflint chert. Science 147:563–577 among astrobiologists for its excellent preserva-
Fralick P, Barrett TJ (1995) Depositional controls on iron tion in stromatolitic and non-stromatolitic cherts
formation associations in Canada. Spec Publ Int Assoc and for its abundance and diversity of filamentous,
Sedimentol 22:137–156 coccoidal fossils and problematic forms that are
House CH, Schopf JW, McKeegan KD, Coath CD, Harri-
son TM, Stetter KO (2000) Carbon isotopic composi- reported only in iron formations between 2.1 and
tion of individual precambrian microfossils. Geology 1.7 Ga.
28:707–710
Knoll AH (2003) Life on a young planet: the first three
billion years of evolution on earth. Princeton Univer-
sity Press, New Jersey, p 304 Overview
Planavsky N, Rouxel O, Bekker A, Shapiro R, Fralick P,
Knudsen A (2009) Iron-oxidizing microbial ecosys- The Paleoproterozoic (2.5–1.6 Ga) era is a crucial
tems thrived in late paleoproterozoic redox-stratified and unique time of changing ocean and atmo-
oceans. Earth Planet Sci Lett 286:230–242
Poulton SW, Fralick PW, Canfield DE (2004) The transi- sphere’s chemistries. During this period, the
tion to a sulphidic ocean 1.84 billion years ago. Nature atmosphere and surface ocean waters were likely
431:173–177 moderately oxygenated (1–10 % PAL; present
Slack JF, Grenne T, Bekker A, Rouxel OJ, Lindberg PA atmospheric level of oxygen). Traces of this
(2007) Suboxic deep seawater in the late paleoproterozoic:
evidence from hematitic chert and iron formation related changing redox conditions are globally detected
to seafloor-hydrothermal sulfide deposits, central Arizona, in the geological record since the GOE (Great
U.S.A. Earth Planet Sci Lett 255:243–256 Oxidation Event, around 2.4 Ga), but local oxy-
Strauss H, Moore TB (1992) Abundances and isotopic com- genation probably occurred earlier. The deeper
positions of carbon and sulfur species in whole rock and
kerogen samples. In: Schopf JW, Klein C (eds) Protero- bottom waters might have been euxinic (anoxic
zoic biosphere. Cambridge University Press, Cambridge and sulfidic; Canfield 1998) or only anoxic or
dysoxic (i.e., having low oxygen levels between
anoxic and hypoxic levels) (Johnston et al. 2009),
with a transient return to a ferruginous ocean
Gunflint Microbiota possibly around Gunflint times, at 1.9 Ga (see
review in Lyons et al. 2009).
Emmanuelle J. Javaux The Paleoproterozoic rocks host the oldest
Palaeobiogeology-Palaeobotany- ▶ microfossils diagnostic of modern
Palaeopalynology, Geology Department, ▶ cyanobacteria in carbonate formations, for
Université de Liège, Liège, Belgium example, the Belcher Fm, Canada, and the
Franceville Fm, Gabon (Amard and Bertrand-
Safarti 1997; Hofmann 1976; review in Knoll
Keywords and Golubic 1992), contemporaneous of the Gun-
flint Fm, and the oldest microfossils of stem-
Gunflint formation; Microfossils; Paleopro- group eukaryotes, in 1.8 Ga shales from China
terozoic; Stromatolites; Traces of life and 1.65 Ga shales from Australia (reviews in
Gunflint Microbiota 1027
Knoll 2003; Knoll et al. 2006). Diverse microfos- umbellata), filaments (Gunflintia), and budding
sils of unknown biological affinities are reported tubular forms (Archaeorestis). Gunflintia fila-
from iron formations around the world, including ments are coated by iron oxides and have been
the Gunflint Fm. The Paleoproterozoic record of compared to modern iron bacteria Leptothrix
biological activities also includes ▶ stromatolites (Knoll and Simonson 1981), and geochemical
and ▶ biomarkers (discussed elsewhere in this analyses have evidenced the activities of
volume), macroscopic carbonaceous compres- Fe-oxidizing bacteria (Planavsky et al. 2009).
sions and possible pyritic fossils, mat rip-ups, Gunflintia and Huroniospora can also be pre-
and putative fossil or trail impressions. served by pyrite, some fossils showing the pres-
ence of small hollow, sometimes C-filled vesicles
Gunflint Microbiota interpreted as cells of heterotrophs degrading
The stromatolitic and nonstromatolitic cherts of them (Wacey et al. 2013). In situ sulfur isotope
the Gunflint Formation have been first investi- analyses of individual microfossils showed that
gated in the classical work of Barghoorn and pyritization resulted from the activity of sulfate- G
Tyler (1965), Awramik and Barghorn (1977), reducing bacteria within anoxic, sulfate-poor
Cloud (1965), and Strother and Tobin (1987). sediment pore waters (Wacey et al. 2013). Car-
The stromatolites have variable chemical compo- bon isotopic analyses performed on individual
sition (iron rich, siliceous, calcitic) and morphol- Gunflint microfossils (Gunflintia and
ogies (from small – sometimes Huroniospora) (House et al. 2000) showed d13C
branching – centimeter-scale columns to large values of 31 ‰ to 37 ‰ consistent with car-
domes up to 1 m in diameter) (Planavsky bon fixation via the Calvin cycle, except for two
et al. 2009). The assemblage includes prokaryotic badly preserved specimens of Hurionospora
taxa difficult to relate to extant organisms. No which had extremely 13C-depleted compositions
eukaryotes are unambiguously recognized, down to 45 ‰. Most microfossils do not show a
although some taxa found in nonstromatolitic preferred orientation and are probably plank-
cherts like Eosphaera (vesicles surrounded by a tonic, but Gunflintia filaments are sometimes
layer of smaller coccoids in an envelope) and densely intertwined and coat convex surfaces,
Leptotrichos (5–30 mm solitary coccoids) could suggesting benthic lifestyle (Hofmann 1969;
be prokaryotic or eukaryotic (Knoll 1996). In the Strother and Tobin 1987).
stromatolitic cherts (Fig. 1), the assemblage con- Similar associations are known in the coeval
tains star-like forms (Eoastrion), coccoids Franceville biota, Gabon (Amard and Bertrand-
(Hurioniospora), problematic forms (Kakabekia Sarfati 1997); the Balbirini Dolomite, Australia
Gunflint Microbiota, Fig. 1 Gunflint microfossils in one star-like specimen of Eoastrion (upper left corner),
thin section through stromatolitic chert (Pictures courtesy filamentous sheath Gunflintia, coccoidal Huroniospora
of K. Lepot), showing in (a) thin filamentous sheaths (bottom right corner), and two large tapering, cellular
Gunflintia and coccoids; (b) double-walled coccoids; (c) filaments of possible cyanobacteria
1028 Gunflint Microbiota
(Oehler 1977); the 1.8 Ga Duck Creek Dolomite, Gunflint Paleoenvironmental Conditions
Western Australia (Knoll and Barghoorn 1976; Geochemical proxies combined with paleonto-
Knoll et al. 1988); the Tyler Fm, Michigan logical investigations may help to constrain the
(Cloud and Morrison 1980); the 1.7 Ga Frere metabolisms and biological affinities of the fila-
Fm, Australia (Walter et al. 1976); and in other ments and isolated cells of the Gunflint
coeval iron formations and subtidal carbonates Fm. ▶ Iron isotopes and REE (rare earth element)
worldwide. These assemblages were widely dis- analyses on hematite-rich stromatolites of the
tributed in shallow habitats where iron-rich deep Gunflint Fm permitted to suggest the presence
waters mixed with oxygenated surface waters, of a shallow redoxcline in the ocean, separating
before the late Paleoproterozoic expansion of fully oxygenated shallow waters from iron-rich
sulfidic subsurface waters. Recent investigations deep waters (Planavsky et al. 2009). In the
of the 1.9 Ga Gunflint Formation (Planavsky redoxcline, microaerophilic chemolithotrophic
et al. 2009) and another slightly younger iron microbial ecosystems could proliferate.
formation, the 1.8 Ga Duck Creek Formation, Cyanobacteria lived in the surface-oxygenated
Western Australia (Wilson et al. 2010), con- waters, as suggested by paleontological investi-
firmed previous micropaleontological investiga- gations (review in Knoll and Golubic 1992) and
tions and provided geochemical evidence C-isotope analyses (Strauss and Moore 1992),
supporting the presence of iron-oxidizing bacte- and provided dissolved oxygen used by chemo-
ria in these early ecosystems. autotrophic Fe-oxidizing bacteria just below.
In most Proterozoic microbial mat assem- This new model could explain the presence of
blages, empty filamentous sheaths (e.g., species cyanobacteria microfossils in some Paleopro-
of the genera Oscilliatoriopsis, Siphonophycus, terozoic carbonate successions and the presence
Eomycetopsis, Tenuofilum, Taeniatum, of iron-coated filaments and other unusual micro-
Gunflintia) are abundant and range from fossils in coeval iron-rich formations.
>10 mm (or >100 mm) to <1 mm in diameter In shallow habitats of the Gunflint Formation,
(Knoll and Golubic 1992). Because of their iron-rich deep waters mixed locally with oxygen-
nondiagnostic morphology, their biological affin- ated surface waters (Planavsky et al. 2009).
ity is difficult to determine in the absence of A recent study based on S-isotopes analyses,
paleoenvironmental interpretations or fossilized and tectonic and sedimentologic models, sug-
behavior. The large sheaths are possibly attrib- gests that the shift to sulfidic conditions in the
uted to cyanobacteria, especially when they occur Paleoproterozoic Animikie Basin (where the
in shallow-water sediments deposited in the pho- Gunflint Fm was deposited) approximately coin-
tic zone and when their distribution shows cided with the onset of restricted circulation.
phototactism (movement induced by a rapid Therefore, the basin may record an evolving
light flashing). They could also represent basin with restricted water circulation and thus
Beggiatoa-like sulfur-oxidizing bacteria forming may not reflect the open-ocean structure and
mats at the sediment-water interface with steep composition during the global onset of sulfidic
redox gradient. The narrow sheaths could also conditions (Pufahl et al. 2010).
represent the remains of Chloroflexus-type pho- Further detailed studies of contemporaneous
tosynthetic bacteria or of Leptothrix-type iron iron formations are required to clarify the global
bacteria (as for Gunflintia minuta, which grew extent of the chemical stratification, changing
in an iron-rich stromatolitic environment of the from a redoxcline between oxygenated surface
Gunflint Formation) (Knoll and Golubic 1992). waters and iron-rich deep waters to a redoxcline
Similar uncertainty applies to colonies of micro- between oxygenated surface waters and sulfidic
fossils such as the possibly heterotrophic deep waters in the late Paleoproterozoic when
rod-shaped Eosynechococcus or the coccoidal iron formations disappear, and the possible impli-
Myxococcoides, in the absence of paleoenvir- cations on ecosystem evolution. The occurrence
onmental information (Knoll and Golubic 1992). of ferruginous waters and iron formation in
Gunflint Microbiota 1029
younger Paleoproterozoic successions (e.g., Barghoorn ES, Tyler SA (1965) Microorganisms from the
1.8 Ga Duck Creek Fm, Western Australia) may Gunflint chert. Science 147:563–577
Canfield DE (1998) A new model for Proterozoic ocean
suggest that the Late Paleoproterozoic expansion chemistry. Nature 396:450–453
of sulfidic subsurface waters was globally asyn- Cloud PE (1965) Significance of Gunflint (precambrian)
chronous (Wilson et al. 2010). microflora. Science 148:27–35
Cloud PE, Morrison K (1980) New microbial fossils from
2 Gyr old rocks in northern Michigan. Geomicrobiol
J 2:161–178
Future Directions Fralick P, Davis DW, Kissin SA (2002) The age of the
Gunflint formation, Ontario, Canada: single zircon
The late Paleoproterozoic represents a unique U-Pb age determinations from reworked volcanic
ash. Can J Earth Sci 39:1085–1091
period in the evolution of the redox conditions Hofmann H (1969) Stromatolites from the Proterozoic
of the atmosphere and ocean, with locally strati- Animikie and Sibley groups. Geological Survey of
fied waters and the development of a redoxcline. Canada Paper, Geological Survey of Canada. 68 p
The Gunflint Formation recorded life and fossil- Hofmann HJ (1976) Precambrian microflora, Belcher G
Islands, Canada: significance and
ization processes in iron-rich oxygen-poor condi- systematic. J Palaeontol 50:1040–1073
tions. Study of past extremophile life and iron- House CH, Schopf JW, McKeegan KD, Coath CD, Harri-
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interdisciplinary investigations, combining tion of individual Precambrian microfossils. Geology
28:707–710
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ciples and applications, vol 1. American Association
of Stratigraphic Palynologists Foundation, Dallas,
pp 51–80
Knoll AH (2003) Life on a young planet: the first three
See Also billion years of evolution on earth. Princeton Univer-
sity Press, New Jersey, p 304
▶ Banded Iron Formation Knoll AH, Barghoorn ES (1976) Gunflint-type microbiota
▶ Biomarkers from Duck Creek Dolomite, Western-Australia. Orig
Life Evol Biosph 7:417–423
▶ Biomarker, Isotopic Knoll AH, Golubic S (1992) Living and Proterozoic
▶ Biomineralization cyanobacteria. In: Schidlowski M et al (eds)
▶ Carbon Isotopes as a Geochemical Tracer Early organic evolution: implications for
▶ Cyanobacteria mineral and energy resources. Springer, Berlin,
pp 450–462
▶ Fossilization, Process of Knoll AH, Simonson B (1981) Early Proterozoic micro-
▶ Gunflint Formation fossils and penecontemporaneous quartz cementation
▶ Iron Isotopes in the sokoman iron formation, Canada. Science
▶ Microfossils 211:478–480
Knoll AH, Strother PK, Rossi S (1988) Distribution and
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▶ Stromatolites Duck Creek Dolomite, Western-Australia. Precam-
Knoll AH, Javaux EJ, Hewitt D, Cohen P (2006) Eukary-
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B 361:1023–1038
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My old cherty stromatolites of the Franceville groups, Oehler DZ (1977) Microflora of the middle Proterozoic
Gabon. Precambrian Res 81:197–221 Balbirini Dolomite (McArthur Group) of Australia.
Awramik SM, Barghorn ES (1977) The Gunflint Alcheringa 2:269–309
microbiota. Precambrian Res 5:121–142
1030 Gyr
Planavsky N, Rouxel O, Bekker A, Shapiro R, Fralick P, Wacey D, McLoughin N, Kilburn MR, Saunders M, Cliff
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tems thrived in late Paleoproterozoic redox-stratified analysis of pyritized microfossils reveals differential
oceans. Earth Planet Sci Lett 286:230–242 heterotrophic consumption in the 1.9-Ga Gunflint
Poulton SW, Fralick PW, Canfield DE (2004) The transi- chert. Proc Natl Acad Sci U S A 110:8020–8024
tion to a sulphidic ocean 1.84 billion years ago. Nature Walter MR, Goode ADT, Hall WDM (1976) Microfossils
431:173–177 from a newly discovered Precambrian stromatolitic iron
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terozoic Animikie Basin record the sulfidic ocean tran- Wilson JP et al (2010) Geobiology of the late Paleopro-
sition? Geology 38:659–662 terozoic Duck Creek Formation, Western Australia.
Strauss H, Moore TB (1992) Abundances and isotopic Precambrian Res 179:135–149
compositions of carbon and sulfur species in whole
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C (eds) The Proterozoic biosphere: a multidisciplinary
study. Cambridge University Press, Cambridge
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huroniospora barghoorn-implications for paleoecol-
ogy of the Gunflint Microbiota. Precambrian Res
36:323–333 ▶ Ga
H
H Bond H2O
▶ Hydrogen Bond ▶ Water in the Solar System

▶ Water in the Universe
▶ Water, Formation and Photodissociation
H+ Region ▶ Water, Solvent of Life
▶ Water, Vibrational and Rotational Transitions
▶ HII Region
+ H2O+
H2Cl
▶ Water, Related Interstellar Radicals and Ions
H2CS H2S
▶ Thioformaldehyde ▶ Sulfur Hydrides in the Interstellar Medium
H2NCN H3O+
▶ Cyanamide ▶ Water, Related Interstellar Radicals and Ions

DOI 10.1007/978-3-662-44185-5
1032 Habitability of the Solar System
liquid water has to be present on a planet to

Habitability of the Solar System support life. ▶ Mars is the next step in the search
for life. Observations by orbiters, landers, and
Antigona Segura rovers indicate that this planet had a wet and
Instituto de Ciencias Nucleares, Universidad warm past, with similar conditions to those that
Nacional Autónoma de México, Mexico, DF, Earth had when life appeared on our planet.
Mexico Further exploration of the Solar System has
shown that there are other interesting places to
search for life, e.g., on one of Jupiter’s satellites,
Keywords ▶ Europa.
Earth; Exoplanets; Habitability; Jupiter satellites;

Life; Mars; Solar System Overview
By comparing Earth with other planets and sat-

ellites of our Solar System, we learn about how
Definition
planets work and what makes our planet habit-
able. In all cases, the support of liquid water on
Habitability for the Solar System refers to places
the surface, underground, or on microenviron-
that may be suitable for carbon-based life. Liquid
ments is relevant for habitability in general (e.g.,
water is the first potential indicator of habitability
Jones and Lineweaver 2010). For remote detec-
on the Solar System bodies (planets and satel-
tion of life, it is required that life generates a
lites). The study of the conditions of these bodies
global signature, that is, a change in the atmo-
is basic to understand the difference between a
sphere or on the surface that can be detected with
habitable planet and one that is not suitable
space-borne telescopes (see, e.g., Segura and
for life.
Kaltenegger 2009).
After Miller (1953) tested the idea of Urey The study of Earth as a planet extends from the
(1952) about the synthesis of prebiotic mole- conditions that promote the emergence of life to
cules from inorganic compounds such as meth- the interaction of geological processes with life.
ane, water, and hydrogen, it was clear that The geologic history of Earth is studied using
abiotic matter could form complex molecules ancient rocks that keep the record of the ancient
that may eventually evolve into the first cells. atmospheric and land processes. Planets and
Organic matter has been found in comets, mete- satellites of the Solar System are the nearest
orites, and interplanetary dust and gas, which experimental bench to explore how planets
pointed out these objects as possible sources of evolve. The study of planetary evolution
the chemical compounds that originated life on starts with understanding the initial conditions
Earth (Brack 1999 and references therein). The that formed planets. Comets, asteroids, and
origin of life still is a matter of study, but we interplanetary dust provide the clues for the
have explored some of the physical and chemical initial composition and processes that occurred
conditions needed for chemical evolution and in the disk where planets formed. Remote obser-
the processes that led to the first common ances- vation of planetary atmospheres and spacecraft
tor. Liquid water seems to be one critical ingre- exploration show the possibilities of environ-
dient for life; therefore, we assume here that ments that planetary bodies can present.
Habitability of the Solar System 1033
Key Research Findings that there may be habitats for living organisms
on the giant planets (Sagan and Salpeter 1976).
Liquid water has been suggested as the best sol- These planets have reducing atmospheres (with
vent for life to emerge and evolve. Water is an large amounts of H2- and H-bearing com-
abundant compound in our Galaxy; it can be pounds), which allows the formation of organic
found in a wide range of environments, from compounds but inhibits reactions to form more
cold dense molecular clouds to hot stellar atmo- complex molecules. Also, the high-speed winds
spheres (see, e.g., Cernicharo and Crovisier (>200 km/s), extreme pressures, and tempera-
2005). Water is liquid at a large range of temper- tures of these giants make their environments
atures and pressures, and, as a solvent, water has too harsh for known organisms (e.g., Atreya
the characteristic of strong polar-nonpolar inter- 1986). Other potentially habitable environ-
actions with certain organic substances. This ments, like extrasolar giant planets’ moons
dichotomy is essential for maintaining stable bio- (e.g., Europa or Titan-like environments) called
molecular and cellular structures (DesMarais the cryo-ecosphere (see, e.g., Peña-Cabrera and
et al. 2002). Polar groups are parts of molecules Durand-Manterola 2004), are not discussed here
with charges that can interact with other charged because we do not have enough information on H
molecules, like water. Nonpolar groups have no their potential habitability.
charge and thus can associate with other nonpolar That leaves us with the terrestrial planets.
molecules, like oil. Mercury is a small rocky planet similar to our
To keep water liquid on its surface, a planet Moon with virtually no atmosphere or water.
needs to maintain a certain temperature and Venus and Mars demonstrate the limits of plan-
pressure range. This leads to the second charac- etary habitability. Venus has a surface pressure
teristic of planets with habitable surface condi- of 92 bars and a surface temperature of 482 C,
tions: an atmosphere. Besides producing a too extreme to sustain known life even though
surface pressure, the atmosphere can stabilize some work suggests possible habitats for life in
the surface temperature of the planet through the clouds (Morowitz 1967; Sagan 1967;
climate feedbacks such as the “▶ greenhouse Seckbach and Libby 1971; Cockell 1999;
effect.” This effect is caused by compounds Schulze-Makuch and Irwin 2002; Schulze-
that are very efficient absorbers in the infrared Makuch et al. 2004). Venus has an atmosphere
but not in the visible. The visible light of the but no surface water. The “floating habitable
parent star reaches and heats the planetary sur- zone” is the region in the Venus middle atmo-
face and is reemitted in the infrared, where part sphere where pressure and temperature condi-
of the energy is absorbed by the atmospheric tions favor the existence of water droplets. If
greenhouse gases, increasing the temperature Venus had a similar water reservoir as Earth, it
of the planet. was converted to vapor due to the high surface
For a planet to have an atmosphere, it has to temperatures of the planet. It remains an open
accrete enough volatiles during its formation question whether Venus lost its water on a “run-
and maintain that atmosphere. For Jovian away greenhouse” or a “moist greenhouse”
planets, this is an easy process, according to the (Kasting et al. 1984; Kasting 1988).
most accepted hypothesis of giant planet forma- Numerous space missions have explored Mars
tion. These planets are formed in a zone of a and its geological history. About 4 billion years
▶ protoplanetary disk that is cold enough for ago, Mars had an atmosphere thick enough to
ices to condense and build a massive planetary maintain liquid water on the Martian surface
core, after which the self-gravity of the planet is (e.g., McKay and Stoker 1989), which led to the
able to retain large amounts of gas. None of the hypothesis of life on early Mars. But today, Mars
gaseous planets in our Solar System show signs is a dry, frozen desert that cannot likely sustain
of life, even though some authors have proposed life on its surface. The Martian atmosphere was
1034 Habitability of the Solar System
partly lost and became too thin to warm the plan- accrete gaseous atmospheres more efficiently
etary surface. The first half billion years of the and would retain for longer their primary atmo-
Solar System were characterized by violent spheres, whereas smaller planets could lose their
impacts; planets were subjected to what is called atmospheres after geological periods of time.
the ▶ Late Heavy Bombardment. During this The presence of a large moon that stabilized
epoch, the leftovers of the building blocks of the obliquity of our planet (Laskar et al. 1993),
planetary formation (asteroids and comets) had as well as the availability of life-forming ele-
frequent collisions with the planets. For Mars, the ments like nitrogen and carbon (Gaidos
result was supposedly catastrophic for its atmo- et al. 2005 and references therein), could further
sphere. Large impactors evaporated the Martian constrain habitability for advanced life.
atmosphere, and the low gravity of the planet was Compared to other terrestrial planets in our
not able to retain the gas in the hot plumes created Solar System, Earth is the perfect place to live;
by those impactors (Melosh and Vickery 1989). it has liquid water on its surface, an atmosphere
In about 1 billion years, Mars lost most of its that keeps it warm, and the right mass to maintain
atmosphere. tectonics.
Plate tectonics could have replenished the
Martian atmosphere. Water and CO2 react with
a planet’s crust forming carbonates; some parts
of the crust are subducted and melted when they See Also
reach the mantle. Volatile compounds like H2O
and CO2 are released into the atmosphere
through volcanism. Plate tectonics results in ▶ Biomarkers Atmospheric, Evolution Over
Geological Time
the release of the remnant heat of the core from
▶ Biomarkers, Spectral
planet formation. If the planet is small, the heat
▶ Europa
will be released faster, and the planet has a short
▶ Giant Planets
period of volcanic activity but not necessarily
plate tectonics. This seems to have been the case
for Mars. Note that the size of the planet does not
▶ Habitable Zone
guarantee active plate tectonics; see, for exam-
▶ Late Heavy Bombardment
ple, Venus.
▶ Mars
Future Directions ▶ Super-Earths
Planetary size is crucial for its habitability,

assuming plate tectonics is essential. In our
Solar System, no planets exist with masses
between Earth (1 M) and Uranus (14.5 M). Atreya SK (1986) Atmosphere and ionospheres of the
The possible characteristics and habitability of outer planets and their satellites. Springer, Berlin
such planets have to be derived from planet Brack A (1999) Life in the solar system. Adv Space Res
24:417–433
formation and geologic evolution models of ter-
Catanzarite J, Shao M, Tanner A, Unwin S, Yu J (2006)
restrial planets. The proposed mass limits for a Astrometric detection of terrestrial planets in the hab-
habitable planet are 1 M (e.g., Turnbull and itable zones of nearby stars with SIM PlanetQuest.
Tarter 2003; Catanzarite et al. 2006) to 10 M Publ Astron Soc Pac 118:1319–1339
Cernicharo J, Crovisier J (2005) Water in space: the water
(e.g., Catanzarite et al. 2006). However, new
world of ISO. Space Sci Rev 119:29–69
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DesMarais DJ et al (2002) Remote sensing of planetary Turnbull MC, Tarter JC (2003) Target selection for
properties and biosignatures on extrasolar terrestrial SETI. I. A catalog of nearby habitable stellar systems.
planets. Astrobiology 2:153 Astrophys J Suppl Ser 145:181–198
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the principle of plentitude: a review of terrestrial Academy of Sciences of the United States of America
planet habitability. Astrobiology 5:100–126 38(4):351–363
Ikoma M, Genda H (2006) Constraints on the mass of a
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Jones EG, Lineweaver CH (2010) To what extent does
terrestrial life “follow the water”? Astrobiology
10(3):349–361 Habitability on Mars
Kasting JF (1988) Runaway and moist greenhouse atmo-
spheres and the evolution of earth and Venus. Icarus Alessandro Airo1, Ernst Hauber2 and Stephan
74:472–494
van Gasselt3
Kasting JF, Pollack JB, Ackerman TP (1984) Response of 1
earth’s atmosphere to increases in solar flux and impli- Institut f€ur Geologische Wissenschaften
cations for loss of water from Venus. Icarus Tektonik und Sedimentäre Geologie, Freie
57:335–355
H
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McKay CP, Stoker CR (1989) The early environment and Deutsches Zentrum f€ur Luft- und Raumfahrt
its evolution on Mars: implications for life. Rev (DLR) e.V., Institut f€ur Planetenforschung,
Geophys 27:189–214 Berlin, Germany
Melosh HJ, Vickery AM (1989) Impact erosion of the 3
Planetary Sciences and Remote Sensing,
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Miller SL (1953) The production of amino acids under Institute of Geological Sciences, Freie
possible primitive Earth conditions. Science Universität Berlin, Berlin, Germany
117:528–529
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215:1259–1260
Oparin AI (1924) Proikhozndenie Zhizni. Izd., Definition
Moskowski Rabochi
Peña-Cabrera GVY, Durand-Manterola HJ (2004) Possi- Habitability has been defined as “the potential of
ble biotic distribution in our galaxy. Adv Space Res
an environment (past or present) to support life of
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Sagan C (1967) Life on the surface of Venus? Nature any kind” (Steele et al. 2005). Consequently, the
216:1198–1199 concept of habitability depends on the conditions
Sagan C, Salpeter EE (1976) Particles, environments, and life requires, which commonly are described as
possible ecologies in the Jovian atmosphere.
complex chemistry taking place within a liquid
Astrophys J Suppl Ser 32:737–755
Schulze-Makuch D, Irwin LN (2002) Reassessing the solvent and with access to energy. It is reasonable
possibility of life on Venus: proposal for an astrobiol- to assume that putative life on Mars would be
ogy mission. Astrobiology 2:197–202 physically implemented on the basis of organic
Schulze-Makuch D, Grinspoon DH, Abbas O, Irwin LN,
chemistry and liquid water (Benner et al. 2004).
Bullock MA (2004) A sulfur-based survival strategy
for putative phototrophic life in the Venusian atmo- Beyond the conceptual requirements for hab-
sphere. Astrobiology 4:11–18 itability (water, organics, energy), life can only
Seckbach J, Libby WF (1971) Vegetative life on Venus or exist in physicochemical settings (e.g., tempera-
investigations with algae which grow under pure CO2
ture, radiation, pH) that do not cause a lethal
in hot acid media and at elevated pressures. In:
Sagan C, Owen TC, Smith HJ (eds) Planetary atmo- degradation of the organisms. Consequently,
spheres, international astronomical union. Symposium habitability greatly depends on the type of bio-
no. 40, Dordrecht, Reidel, p 62 chemistry, which makes it challenging to predict
Segura A, Kaltenegger L (2009) In: Basiuk VA, Navarro-
the physicochemical boundaries of habitability
González R (eds) Astrobiology: emergence, search
and detection of life. American Scientific Publishers, with respect to alien life of unknown molecular
2010 composition.
1036 Habitability on Mars
Although this difficulty cannot be overcome, Mars Today

the discovery of extremophiles over the past few
decades has greatly expanded our knowledge of Although Mars might have been habitable at the
the range of habitable environments for “life as surface in the past, today the Martian surface
we know it”. Based on these known boundaries of conditions (low temperatures and pressures)
life on Earth, it can be conservatively estimated usually do not allow liquid water to be stable.
that habitable environments on Mars are those Only at very low-lying areas (higher atmo-
where liquid water is stable at temperatures spheric pressure) and/or if the water is very
below 120 C. saline (reduced melting point temperature) can
water remain liquid at the surface today. Local
melting of ground ice and the formation of cold
Early Mars brines are hypothesized to be responsible for the
seasonal appearance of recurring slope lineae
Among the first to scientifically discuss the puta- (McEwen et al. 2011). However, such environ-
tive habitability of Mars was Alfred Russel Wal- ments would only be habitable for halophilic
lace (Wallace 1907). Although Wallace largely and psychrophilic organisms that are known
argued against Mars being habitable, high hopes to thrive in arctic environments on Earth. Alter-
for this not being the case persisted until the natively, in the event of a large impact into
1970s when the Viking landers reported Mars to water-ice-bearing terrain, crater lakes could
be a cold desert. form and be considered to represent transitional
Since then, Mars exploration efforts were habitable surface environments (McKay and
guided by the slogan “follow the water” and Davis 1991).
yielded a profound understanding of Mars’ watery In contrast to surface environments, it could
past and the current water-ice-rich cryosphere well be that putative subsurface aquifers are the
(e.g., Lasue et al. 2013). As a result of this largest, most stable, and longest-lasting habit-
successful search, it symbolically transitioned to able environments on Mars. Furthermore, it can
“follow the habitability.” The recent discovery of be argued that if microbial life did exist on Early
clay-rich sediments at Gale Crater by the Curiosity Mars, it is more likely that it has adapted to the
rover suggests that these deposits formed within a deep subsurface conditions, as life has done on
shallow-water environment of neutral pH and Earth, than it having gone extinct entirely.
therefore would indicate the former existence of
a habitable lacustrine environment on Early Mars
(Grotzinger et al. 2014).
However, the frequency, duration, and chem- See Also
istry of such habitable paleoenvironments
remain largely unknown, and the degree of sim- ▶ Aquifer (Mars)
ilarity between Early Mars’ and Earth’s habit- ▶ Crater Lakes (Mars)
able environments continues to be debated. ▶ Extremophiles
Furthermore, the length of time needed and ▶ Mars
exact physicochemical conditions required for
an origin of Earth-like life remain in the realm
of speculation. Therefore, any estimate on the References and Further Reading
suitability of Martian paleoenvironments for an
origin of life is highly speculative as well. How- Benner SA et al (2004) Is there a common chemical model
ever, even if life never originated on Mars itself, for life in the universe? Curr Opin Chem Biol
8:672–689
it could have been transferred from Earth to
Grotzinger JP et al (2014) A habitable fluvio-lacustrine
Mars through the process of ▶ panspermia and environment at yellow knife bay, gale crater, Mars.
continued to thrive on Mars until today. Science 343:386–387
Habitability, Effects of Stellar Irradiation 1037
Lasue J, Mangold N, Hauber E, Clifford S, Feldman W, ▶ Habitable Planet, Characterization

Gasnault O, Grima C, Maurice S, Mousis O (2013) ▶ Habitable Zone
Quantitative assessment of the Martian hydrosphere.
Space Sci Rev 174:155–212 ▶ Periastron
McEwen AS, Ojha L, Dundas CM, Mattson SS, Byrne S,
Wray JJ, Cull SC, Murchie SL, Thomas N, Gulick VC
(2011) Seasonal flows on warm Martian slopes. Sci- References and Further Reading
ence 333:740–743
McKay CP, Davis WL (1991) Duration of liquid water Dressing C et al (2010) Astrophys J 721(2):1295–1307
habitats on early Mars. Icarus 90:214–221 Spiegel DS et al (2010) Generalized milankovitch cycles
Steele A. et al. (2005) The astrobiology field laboratory. and long-term climatic habitability. Astrophys
Unpublished white paper, 72 p, posted Dec., 2005 by J 721(2):1308–1318
the Mars Exploration Program Analysis Group Williams DM, Pollard D (2002) Earth-like worlds on
(MEPAG) at http://mepag.jpl.nasa.gov/reports/index. eccentric orbits: excursions beyond the habitable
html zone. Int J Astrobiol 1:61
Wallace AR (1907) Is Mars habitable? MacMillan and
Co., Limited, London
H
Habitability, Effects of Stellar
Irradiation
Habitability, Effect of Eccentricity
Lisa Kaltenegger
Lisa Kaltenegger Cornell University, Ithaca, NY, USA
Keywords
Definition
Biomarkers; Earth geological evolution of;
Variations in the ▶ eccentricity of a planet’s orbit Habitability; Biomarkers remote detectability
can lead to climate changes, including possible thereof
phase transitions of liquids on a planet’s surface
and therefore different climate feedback mecha-
nisms due to ▶ albedo changes. A more eccentric Definition
orbit both intensifies the ratio of stellar irradiation
at ▶ periastron to that at apoastron and increases Stellar irradiation refers in this context to the
the annually averaged irradiation, proportional to wavelength-dependent incoming flux of photons
1=2
ð1 e2 Þ , where e is eccentricity. into the planetary atmosphere from the central
Periodic oscillations of eccentricity cause star. This in turn will influence e.g. the planet’s
oscillations of the total amount of starlight inci- atmospheric photochemistry, climate, and poten-
dent on a planet in each annual cycle. These tial biomarkers.
oscillations depend on gravitational perturbations
from other companion objects.
Overview
See Also The range of characteristics of planets is likely

to exceed our experience with the planets and
▶ Albedo satellites in our own Solar System by far.
▶ Atmosphere, Structure Models of planets more massive than our
▶ Eccentricity Earth need to consider the changing atmosphere
▶ Habitability of the Solar System structure, as well as the interior structure of the
1038 Habitability, Effects of Stellar Irradiation
planet (see, e.g., Seager et al. 2007; Valencia See Also

et al. 2006). Earthlike planets orbiting stars of
different spectral type might evolve differently ▶ Atmosphere, Structure
(Selsis 2000; Segura et al. 2003, 2005). Model- ▶ Biomarkers Atmospheric, Evolution Over
ing these influences will help to optimize the Geological Time
design of the proposed instruments to search for ▶ Biomarkers, Spectral
Earthlike planets. The spectral resolution ▶ Habitable Planet, Characterization
required for optimal detection of habitability ▶ Habitable Zone
and biosignatures has to be able to detect those ▶ Spectral Type
features for different stellar types as well as
over the evolution of a planet (▶ Biomarkers,
Spectral; Biomarkers, Atmospheric; ▶ Habit-
able Planet, Characterization). Using a numeri-
cal code that simulates the photochemistry of a Grenfell JL, Stracke B, von Paris P, Patzer B, Titz R,
wide range of planetary atmospheres, several Segura A, Rauer H (2007) The response of atmo-
groups (Selsis 2000; Segura et al. 2003, 2005; spheric chemistry on earthlike planets around F,
Grenfell et al. 2007) have simulated a replica of G and K Stars to small variations in orbital distance.
Planet Space Sci 55:661–671
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F-type star (more massive and hotter than the targets for terrestrial exoplanet searches and
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(2007) Mass-radius relationships for solid exoplanets.
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Habitable Planet, Characterization 1039
Overview
Habitable Planet, Characterization
We discuss how we can read a planet’s spectrum
Lisa Kaltenegger1 and Franck Selsis2,3 to assess its habitability and search for the signa-
1
Cornell University, Ithaca, NY, USA tures of a biosphere. After a decade rich in giant
2
Laboratoire d’Astrophysique de Bordeaux, exoplanet detections, observation techniques
Universite de Bordeaux, Floirac, France have now reached the ability to find planets of
3
CNRS, LAB, Floirac, France less than 10 MEarth (so-called ▶ super-Earths)
that may potentially be habitable. Extrasolar
planet searches have shown an extraordinary
Keywords ability to combine research with astrophysics,
chemistry, biology, and geophysics into a new
Biomarkers; Extrasolar planets; Habitability; and exciting interdisciplinary approach to under-
Habitable zone; Planetary atmospheres; standing our place in the universe.
Spectroscopy
H
Introduction
Definition The current status of exoplanet characterization
shows a surprisingly diverse set of giant planets.
The spectrum of a planet can contain signatures For a subset of these, some properties have been
of atmospheric species, which create its spectral measured or inferred using observations of
fingerprint. The presence and abundance of atmo- the host star, a background star, or the combina-
spheric species, in the context of the properties of tion of stellar and planetary photons (▶ Radial-
the star and the planet, can elucidate the underly- Velocity Planets, ▶ Microlensing Planets,
ing physics and characterize a planetary environ- ▶ Transiting Planets, and ▶ Astrometry). These
ment. Here, we concentrate on characterizing a observations have yielded measurements of
habitable planet and see what features might have planetary mass, orbital elements and (for tran-
a biological origin. sits) the planetary radius, and, during the last few
years, physical and chemical characteristics of
the upper atmosphere of some of the transiting
History planets.
The detection of an Earthlike planet is
Sagan et al. (1993) analyzed a spectrum of the approaching rapidly thanks to radial velocity
Earth taken by the Galileo probe, searching for surveys (▶ HARPS), transit searches (Corot,
signatures of life, and concluded that the large Kepler), and space observatories dedicated to
amount of O2 and the simultaneous presence of their characterization that are already in
traces of CH4 are strongly suggestive of biology. the development phase (▶ James Webb Space
To characterize a planet’s atmosphere and its Telescope), as well as future large ground-
potential habitability, we look for absorption fea- based telescopes and dedicated space-based
tures in the reflection and transmission spectrum missions. Space missions like CoRoT (CNES;
of the planet. On Earth, some atmospheric species Rouan et al. 1998) and Kepler (NASA; Borucki
exhibiting noticeable features in the planet’s et al. 1997) provide statistics on the number,
spectrum result directly or indirectly from bio- size, period, and orbital distance of planets,
logical activity: The main ones are O2, O3, CH4, extending to terrestrial planets on the lower
and N2O. CO2 and H2O are, in addition, impor- mass range end, while future space missions
tant as ▶ greenhouse gases in a planet’s atmo- are designed to characterize their atmospheres.
sphere and potential sources for a high O2 Future space missions have the explicit
concentration from photosynthesis. purpose of detecting other Earthlike worlds,
1040 Habitable Planet, Characterization
Life ? What biota?
H2O H2O What chemistry?

H2O H2O Atmosphere
CO2 CO2
What star? HZ?
O3
CH4 Interior Plate tectonics?
Ocean/land?
What kind Density?
(radius/mass)
Formation Formation models
Habitable Planet, Characterization, Fig. 1 Connection between spectra and characterization of rocky exoplanets
10−6
10−7
10−8
10−9
10−10 3.0
10−11
lλ erg/(cm2 s mm)
Star F2V
10−12 2.5
Flux (W m−2 nm−1)
10−13 2.0
Sun
10−14
K2V Non-active star
10−15 1.5 Teff = 3100K
10−16 J J GJ 643
1.0
10−17 V E
AD Leo
E
10−18 M
V
M 0.5
10−19 Z Z
10−20 0.0
0.1 1 10 100 0 1000 2000 3000
λ (mm) Wavelength (nm)
Habitable Planet, Characterization, Fig. 2 SAO model of our solar system (left) (assumed here to be black bodies
with Earth spectrum shown). Spectra of different host stars (Segura et al. 2005) (right)
analyzing their characteristics, determining the Basic Methodology

composition of their atmospheres, investigating
their capability to sustain life as we know it, and Characterize a Habitable Planet
searching for signs of life. This can set our own A planet is a very faint, small object close to a
planet, the only known habitat, in context with very bright and large object, its parent star. In the
other rocky worlds and expand our statistics of visible part of the spectrum, we observe the star-
3 for extrasolar rocky planets that could poten- light reflected off the planet, while in the infrared
tially support life (▶ Habitability of the Solar (IR), we detect the planet’s own emitted flux. The
System). They have the capacity to investigate Earth-Sun intensity ratio is about 107 in the
the physical properties and composition of a thermal infrared (10 mm) and about 1010 in
broader diversity of planets, to understand the the visible (0.5 mm) (see Fig. 2). The trade-off
formation of planets, and to interpret potential between contrast ratio and design is not discussed
biosignatures (see Fig. 1). here, but leads to several different configurations
for space-based mission concepts. The suggested model fits to spectra of the Earth in three wave-
interferometric systems operate in the mid IR length ranges (Kaltenegger et al. 2007). The data
(6–20 mm) and observe the thermal emission shown in Fig. 3 (left) is the visible Earthshine
emanating from the planet. The coronagraph spectrum (Woolf et al. 2002), (right) is the near-
and occulter concepts detect the reflected light infrared Earthshine spectrum (Turnbull
of a planet and operate in the visible and near et al. 2006), and (bottom) is the thermal infrared
infrared (0.5–1 mm). The viewing geometry spectrum of Earth as measured by a spectrometer
results in different flux contributions of the over- en route to Mars (Christensen and Pearl 1997).
all detected signal from the planet’s bright and The data are shown in black and the SAO model
dark side for the reflected light and the planet’s in red. In each case, the constituent gas spectra in
hot and cold regions for the emitted flux. The a clear atmosphere are shown in the bottom panel,
contrast ratio of hot extrasolar giant planets for reference.
(EGP) to their parent stars’ flux is much smaller Both spectral regions contain the signature of
and can partially be observed with current tele- atmospheric gases that may indicate habitable
scopes. The contrast ratio of a rocky planet to a conditions and, possibly, the presence of a bio-
smaller parent star is much more favorable, mak- sphere: CO2, H2O, O3, CH4, and N2O in the H
ing Earthlike planets around small stars very thermal infrared and H2O, O3, O2, CH4, and
interesting targets for the immediate future. CO2 in the visible to near-infrared in reflected
Our search for signs of life is based on the (Fig. 4 left). The Earth’s spectra are shown in
assumption that extraterrestrial life shares funda- Fig. 4. The presence or absence of these spectral
mental characteristics with life on Earth, in that it features (detected individually or collectively)
requires liquid water as a solvent, has an energy will indicate similarities or differences with the
source, and features a carbon-based chemistry atmospheres of terrestrial planets and their
(see, e.g., Brack 1993; Des Marais et al. 2002). astrobiological potential (see Kaltenegger and
Life on the basis of a different chemistry is not Traub (2009) and Palle et al. (2009) for details
considered here because the vast range of possi- on Earth’s transmission spectrum).
ble life-forms might produce signatures in their
planet’s atmosphere that are so far unknown.
Therefore, we assume that extraterrestrial life is Key Research Findings
similar to life on Earth in its use of the same input
and output gases and that it exists out of thermo- Characterizing Planetary Environments
dynamic equilibrium (Lovelock 1975). “▶ Bio- It is relatively straightforward to remotely ascer-
markers” are used here to mean detectable tain that Earth is a habitable planet, replete with
species, or a set of species, whose presence at oceans, a greenhouse atmosphere, global geo-
significant abundance strongly suggests a biolog- chemical cycles, and life – if one has data with
ical origin (e.g., the simultaneous presence of arbitrarily high signal-to-noise ratio and spatial
CH4 + O2 or CH4 + O3 (Lovelock 1975)). and spectral resolution. The interpretation of
Bioindicators are indicative of biological pro- observations of other planets with limited signal-
cesses but can also be produced abiotically. It is to-noise ratio and spectral resolution, as well as
their abundance and detection along with other absolutely no spatial resolution as envisioned for
atmospheric species and in a certain context (for the first-generation search instruments, will be far
instance, the properties of the star and the planet) more challenging and implies that we need to
that point toward a biological origin. The spec- gather information on the planetary environment
trum of the planet can contain signatures of atmo- to understand what we will see.
spheric species, which creates its spectral The following step-by-step approach can be
fingerprint (▶ Biomarkers Atmospheric, Evolu- taken to set the planetary atmosphere in context.
tion Over Geological Time and ▶ Biomarkers, After detection, we will focus on the main prop-
Spectral). Figure 3 shows observations and erties of the planetary system: its orbital elements
1.2
0.8
Data
1
0.6
Data
0.8 Model
Relative reflectane
0.4 Model
0.6
0.2
0.4
1 0
0.8 O 3 O2 O2 O2 H2O 0.8 CH4
H2O O2 CO2
0.6 0.6
0.4 0.4
0.2 0.2
0 0
0.6 0.8 1 1.2 1.4 1.6 1.8
Wavelength (mm) Wavelength (mm)
10
Data
Model
0
1
CH4
H2O O3 CO2 H 2O
0
5 10 15 20
Wavelength (mm)
Habitable Planet, Characterization, Fig. 3 Observed determined from Earthshine and space, respectively. The
reflectivity spectrum in the visible (Woolf et al. 2002) data is shown in black and the Smithsonian Astrophysical
(top), near-infrared (Turnbull et al. 2006) (middle), and Observatory (SAO) model in red. The reflectivity scale is
emission spectrum in the infrared (Christensen and Pearl arbitrary
1997) (bottom panel) of the integrated Earth, as
as well as the presence of an atmosphere, using have an abiotic explanation of all compounds
the light curve of the planet, or/and a crude esti- seen in the atmosphere of such a planet. If we
mate of the planetary nature from very do not, we can work with the exciting biotic
low-resolution information (three or four wave- hypothesis. O2, O3, and CH4 are good biomarker
length channels). Then a higher-resolution spec- candidates that can be detected by a
trum will be used to identify the compounds of low-resolution (resolution <50) spectrograph.
the planetary atmosphere and to constrain the Note that if the presence of sets of biogenic
temperature and radius of the observed gases such as (O2/O3 + CH4) may imply the
exoplanet. In that context, we can then test if we presence of a massive and active biosphere,
1 Total transit
Relative transmission
0.8
O2 HNO3 H2O
0.6 H2O CH
O3 H2O 4
O3
10−6 λ/Δλ = 100 0.4
CO2 CO2
Earth/Sun flux ratio
H2O 0.2
10−7 CO2 0
blackbody O3 5 10 15 20
−8 (288 K)
Wavelength (mm)
10
O3 60
Effective height (km)

CO2
10−9 O3 O3 H2O CO2 CO2
O3
H2O 40 H2O
10−10 O3
O2 CO2 O2 H2O CH4
20 HNO3 H2O
10−11
CO2
10−12 0
1 10 5 10 15 20
Wavelength (mm) Wavelength (mm) H
Habitable Planet, Characterization, Fig. 4 Synthetic intensity is given as a fraction of solar intensity (left) as
reflection and emission spectra (left) and transmission well as the relative height in the atmosphere. The atmo-
spectra (right) of the Earth from UV to IR shown. The spheric features are indicated
their absence does not imply the absence of life. (regions of transparency) which allow probing
Life existed on Earth before the interplay the surface or certain atmospheric levels. Such
between oxygenic photosynthesis and carbon identification is not trivial. For an Earthlike
cycling produced an oxygen-rich atmosphere. planet, there are some atmospheric windows
that can be used in most of cases, especially
Temperature and Radius of a Planet between 8 and 11 mm as seen in Fig. 4. This
Knowing the temperature and planetary radius is window would, however, become opaque at
crucial for the general understanding of the phys- high H2O partial pressure (e.g., in the inner part
ical and chemical processes occurring on the of the ▶ habitable zone (HZ) where a lot of water
planet (tectonics, hydrogen loss to space). In the- is vaporized) and at high CO2 pressure (e.g., a
ory, spectroscopy can provide some detailed very young Earth or the outer part of the HZ). The
information on the thermal profile of a planetary accuracy of the radius and temperature determi-
atmosphere (▶ Atmosphere, Structure). This, nation will depend on the sensitivity and resolu-
however, requires a spectral resolution and a sen- tion of the spectrum, the precision of the Sun-star
sitivity that are well beyond the performance of a distance, the cloud coverage, and also the distri-
first-generation spacecraft. Here we concentrate bution of brightness temperatures over the plan-
on the initially available observations. etary surface. For transiting planets, the accuracy
One can calculate the stellar energy of the star of the radius of the planet depends on how well
Fstar that is received at the planet’s measured the host star is characterized.
orbital distance. The surface temperature of the The measured IR flux can directly be
planet at this distance depends on its ▶ albedo converted into a brightness temperature that will
and on the greenhouse warming by atmospheric provide information on the temperature of the
compounds. However, with a low-resolution atmospheric layers responsible for the emission.
spectrum of the thermal emission, the mean If the mass of a planet can be measured (by radial
effective temperature and the radius of the planet velocity and/or astrometric observations), an esti-
can be obtained. The ability to associate a surface mate of the radius of the rocky core can be made
temperature to the spectrum relies on the exis- by assuming a bulk composition of the planet
tence and identification of spectral windows which can then be used to convert IR fluxes into
Visible-NIR (reflected light)

1.4 1.4 IR (thermal emission)
oon
for all planets
rM
ry o
rcu
s
1.2 1.2
ar
Me
M
on
<flux>period
<flux>period
+Mo
Earth Earth
flux
flux
Venus
1.0 1.0
0.8 0.8
0.6 0.6
0.0 0.2 0.4 0.6 0.8 1.0 0.0 0.2 0.4 0.6 0.8 1.0
Orbital phase Orbital phase
Habitable Planet, Characterization, Fig. 5 Orbital the visible (left) and thermal infrared (right) (After Selsis
▶ light curve for ▶ blackbody planets in a circular orbit 2002). Flux normalized to the average over a complete
with null obliquities, with and without an atmosphere in period
temperatures. Important phase-related variations Note also that a Venus-like exoplanet would
in the planet’s flux are due to a high day/night exhibit nearly no measurable phase-related vari-
temperature contrast and imply a low ▶ green- ation of its thermal emission, due to the fast
house effect and the absence of a stable liquid rotation of its atmosphere and its strong green-
ocean. Therefore, habitable planets can be distin- house effect, and thus can only be distinguished
guished from airless or Mars-like planets by the through spectroscopy from habitable planets. The
amplitude of the observed variations of Tb (see mean value of Tb estimated over an orbit can be
Fig. 5). used to estimate the ▶ albedo of the planet, A,
The orbital flux variation in the IR can distin- through the balance between the incoming stellar
guish (in the detection phase) planets with and radiation and the outgoing IR emission.
without an atmosphere (see also Selsis 2002; The thermal light curve (i.e., the integrated
Gaidos and Williams 2004). Strong variation of infrared emission measured at different positions
the thermal flux with the phase reveals a strong on the orbit) exhibits smaller variations due to the
difference in temperature between the day and phase (whether the observer sees mainly the day-
night hemisphere of the planet, a consequence side or the nightside) and to the season for a planet
of the absence of a dense atmosphere which with an atmosphere, than does the corresponding
would transport heat between the hemispheres. visible light curve (see Fig. 5). In the visible
In such a case, estimating the radius from the ranges, the reflected flux allows us to measure the
thermal emission is made difficult because most product A R2, where R is the planetary radius
of the received flux comes from the small and hot (a small but reflecting planet appears as bright as a
substellar area. The ability to retrieve the radius big but dark planet). The first generation of optical
would depend on the assumptions that can be instruments will be very far from the angular res-
made on the orbit geometry and the rotation rate olution required to directly measure an exoplanet
of the planet (▶ Habitable Zone, Effect of Tidal radius.
Locking). In most cases, degenerate solutions
will exist. When the mean brightness temperature Potential Biomarkers
is stable along the orbit, the estimated radius is Spectral ▶ biomarkers are discussed in a separate
more reliable. The radius can be measured at entry in this encyclopedia. We simply add here
different points of the orbit and thus for different Fig. 6 on the terrestrial oxygen cycle. Note that
values of brightness temperature Tb, which possible abiotic sources of biomarkers must be
should allow estimating the error made. carefully evaluated.
Atmosphere
1,100,000 Gt-0
2nH2O* + nCO2 + hv 0.13 Gt-0/yr
Respiration Net consumption of O2
0.13 Gt-O/ayr
190 Gt-0/yr Photosynthesis by oxidation of fossil
Net release of O2 190 Gt-0/yr carbon
Photosynthesis after carbon burial
190 Gt-0/yr Biosphere
7,500 Gt-O
nH2O + n(CHO2O) + nO*2
2,800 Gt-C
Respiration/
oxidation 0.05 Gt-C/yr 0.05 Gt-C/yr
190 Gt-0/yr carbon burial Flux of fossil carbon
to the surface
Sediments
(buried organics)
10,000,000 Gt-C
H
Habitable Planet, Characterization, Fig. 6 Oxygen cycle on Earth (Kaltenegger and Selsis 2010); Gt is gigatons
Cryptic Worlds, Surface Features and Cloud surface reflectivity, one could discover conti-
Features nents and seas on an exoplanet by observing the
▶ Clouds reduce the relative depths, full widths, daily variation of the surface albedo in the visible
and equivalent widths of spectral features, weak- (Ford et al. 2001; Palle et al. 2008). On a cloud-
ening the spectral lines in the whole described as free Earth, the diurnal flux variation at visible
wavelength range (Kaltenegger et al. 2007). Earth wavelengths caused by different surface features
has an average of 60 % cloud coverage, which rotating in and out of view could be high, assum-
prevents easy identification of any surface features ing hemispheric inhomogeneity. When the planet
without knowing the cloud distribution. If one is only partially illuminated, a more concentrated
records the planet’s signal with a very high time signal from surface features could be detected as
resolution (a fraction of the rotation period of the they rotate in and out of view on a cloudless
planet) and individually high signal-to-noise ratio planet (William and Gaidos 2008).
(SNR), one could determine the overall contribu- Our knowledge of the reflectivity of different
tion of clouds to the signal (Cowan et al. 2009; surface components on Earth – like desert, ocean,
Palle et al. 2008). During each of these individual and ice – helps in assigning the vegetation red
measurements, one has to collect enough photons edge (VRE) of the Earthshine spectrum to terres-
for a high individual SNR per measurement to be trial vegetation. On Earth around 440 million
able to correlate the measurements to the surface years ago (Pavlov et al. 2003; Schopf 1993), an
features, which precludes this method for first- extensive land plant cover developed, generating
generation missions that will observe a minimum the red chlorophyll edge in the reflection spec-
of several hours to achieve an SNR of 5–10. For trum between 700 and 750 nm. While they effi-
Earth (Cowan et al. 2009; Palle et al. 2008), these ciently absorb visible light, photosynthetic plants
measurements show a correlation to Earth’s sur- have developed strong infrared reflection
face features, because the individual measure- (possibly as a defense against overheating and
ments are time resolved as well as have an chlorophyll degradation) resulting in a steep
individual high SNR, making it a very interesting change in reflectivity around 700 nm, called the
concept for future generations of missions. red edge. The primary molecules that absorb the
Assuming one had a planet with a cloud-free energy and convert it to drive photosynthesis
atmosphere or could distinguish the signal from (H2O and CO2 into sugars and O2) are chloro-
the overall cloud distribution from that due to the phyll A (0.450 mm) and B (0.680 mm). The exact
wavelength and strength of the spectroscopic Summary

“vegetation red edge” (VRE) depend on the Any information we collect on habitability is
plant species and environment. Averaged over a only important in a context that allows us to
spatially unresolved hemisphere of Earth, the interpret what we find. To search for signs of
additional reflectivity of this spectral feature is life, we need to understand how the observed
typically only a few percent (see also (Montanes- atmosphere physically and chemically works.
Rodriguez et al. 2005; Tinetti et al. 2006; Knowledge of the temperature and the planetary
Kaltenegger and Traub 2009). Several groups radius is crucial for the general understanding of
(Arnold et al. 2002; Christensen and Pearl 1997; the physical and chemical processes occurring on
Montanes-Rodriguez et al. 2007; Woolf the planet. These parameters, as well as an indi-
et al. 2002; Turnbull et al. 2006) have measured cation of habitability, can be determined with
the integrated Earth spectrum via the technique of low-resolution spectroscopy and low photon
Earthshine, using sunlight reflected from the flux, as assumed for first-generation space mis-
non-illuminated, or “dark,” side of the Moon. sions. The combination of spectral information in
Earthshine measurements have shown that detec- the visible (starlight reflected off the planet) as
tion of Earth’s VRE is feasible if the resolution is well as in the mid IR (planet’s thermal emission)
high and the cloud coverage is known. On planets allows a confirmation of detection of atmospheric
around other stars, different biota could develop species and a more detailed characterization of
and produce different reflective features (Kiang individual planets and the opportunity to explore
et al. 2007). a wide domain of planet diversity. Being able to
On Earth, photosynthetic organisms are respon- measure the outgoing shortwave and longwave
sible for the production of nearly all of the oxygen radiation as well as their variations along the
in the atmosphere. However, in many regions of orbit, to determine the albedo and identify green-
the Earth and particularly where surface conditions house gases, would in combination allow us to
are extreme, for example, in hot and cold deserts, explore the climate system at work on the
photosynthetic organisms can be driven into and observed worlds, as well as probe planets similar
under substrates where light is still sufficient for to our own for habitable conditions.
photosynthesis. These communities exhibit no
detectable surface spectral signature. The same is
true of the assemblages of photosynthetic organ- Future Directions
isms at more than a few meters depth in water
bodies. These communities are widespread and The results of a first-generation mission will most
dominate local photosynthetic productivity. likely include an amazing scope of diverse planets
There could be such very interesting Earth-analog that will set planet formation and evolution, as well
worlds that could have habitats but not exhibit as our own planet, in an overall context.
biological surface feature in the disk-averaged
spectrum (Cockell et al. 2009).
Earth’s hemispherical integrated vegetation See Also
red edge signature is very weak, but planets
with different rotation rates, obliquities, ▶ Albedo
land-ocean fraction, and continental arrangement ▶ Astrometry
may have lower cloud cover and higher vegetated ▶ Atmosphere, Structure
fraction. Knowing that other pigments exist on ▶ Biomarkers Atmospheric, Evolution Over
Earth and that some minerals can exhibit a similar Geological Time
spectral shape around 750 nm (Seager ▶ Biomarkers, Spectral
et al. 2005), the detection of the red edge of ▶ Blackbody
chlorophyll on exoplanets, despite its extreme ▶ Clouds
interest, will not be unambiguous. ▶ CoRoT Satellite
▶ Greenhouse Effect Ford E, Seager S, Turner EL (2001) Characterization of

▶ Habitability, Effect of Eccentricity extrasolar terrestrial planets from diurnal photometric
variability. Nature 412:885–887
▶ Habitability, Effects of Stellar Irradiation Forget P, Pierehumbert H (1997) Warming early Mars
▶ Habitability of the Solar System with carbon dioxide clouds that scatter infrared radia-
▶ Habitable Zone tion. Science 278:1273–1274
▶ Habitable Zone, Effect of Tidal Locking Gaidos E, Williams DM (2004) Seasonality on terrestrial
extrasolar planets: inferring obliquity and surface con-
▶ HARPS ditions from infrared light curves. New Astron 10:67–72
▶ Kepler Mission Grenfell JL, Stracke B, von Paris P, Patzer B, Titz R,
▶ Lightcurve Segura A, Rauer H (2007) The response of atmo-
▶ Microlensing Planets spheric chemistry on earthlike planets around F,
G and K Stars to small variations in orbital distance.
▶ Radial-Velocity Planets Planet Space Sci 55:661–671
▶ Super-Earths Harrington J, Hansen BM, Luszcz SH, Seager S,
▶ Transiting Planets Deming D, Menou K, Cho JY-K, Richardson LJ
(2006) The phase-dependent infrared brightness of
the extrasolar planet u Andromedae b. Science
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Kalas P, Graham JR, Chiang E, Fitzgerald MP,
H
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Keywords
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In: Foing B, Battrick B (eds) Earth-like planets and Exoplanets; Habitability; Life
Habitable Zone 1049
Definition available. Such metabolisms rely on very limited

sources of energy (compared to stellar light) and
The circumstellar habitable zone (HZ) is defined electron donors (compared to H2O on Earth).
as the annulus around a ▶ main sequence star They mainly catalyze reactions that would occur
where a rocky planet similar to Earth in compo- at a slower rate in purely abiotic conditions, and
sition and mass with an atmosphere can support they are thus not expected to modify a whole
liquid water on its surface. The inner edge is planetary environment in a way detectable
defined (Kasting et al. 1993) as the distance at remotely.
which a planet undergoes runaway greenhouse
conditions vaporizing the whole water reservoir
and, as a second effect, inducing the photodisso- Basic Methodology
ciation of water vapor and the loss of hydrogen
to space. The outer boundary is defined as the The equilibrium temperature Teq of a planet may
distance from the star where a maximum be found by equating the stellar energy flux
▶ greenhouse effect fails to keep the surface of absorbed to the thermal energy that is radiated
the planet above freezing or the distance back to space. The result is H
from the star where CO2 gas starts condensing
(see, e.g., Kasting et al. 1993). The width 1=
and distance of this annulus, using this defini- Teq ¼ ðSð1 AÞ=fsÞ 4 (1)
tion, depend mainly on the stellar luminosity
and spectral energy distribution. A planet in the where A is the Bond ▶ albedo, S the stellar
habitable zone is not necessarily habitable. energy flux, s the Stefan-Boltzmann constant,
Multiple mechanisms exist by which a planet and f a redistribution factor that accounts for the
cannot attain or can lose habitability, for exam- movement of energy around the planetary surface
ple, low mass and resulting atmospheric losses due to rotation (▶ Habitable Zone, Effect of Tidal
or changes in initial composition (e.g., no Locking), atmospheric circulation, etc. The Bond
water). ▶ albedo is 0.29 for Earth and is defined as the
fraction of power at all wavelengths that is
scattered back out into space. If all the incident
History energy is uniformly distributed on the planetary
sphere (e.g., by atmospheric circulation), f = 4;
The habitable zone (HZ) concept was proposed if only the day hemisphere is so heated, f = 2. If
for the first time by Huang (1959, 1960) and has there is no redistribution, f = 1/cos(F) where F
been calculated by several authors after that (see, is the zenith angle. The last case gives good
e.g., Rasool and DeBergh 1970; Hart 1979; results for the surface temperature on the sunlit
Kasting et al. 1993; Kopparapu et al. 2012). hemisphere of airless bodies with known albedo
like the Moon and Mercury. Note that for a planet
with a dense atmosphere, Teq is not necessarily
Overview equal to the physical temperature at the surface,
but may refer to a level in the atmosphere. With
Future remote sensing characterization of plane- albedos of 0.75, 0.3, and 0.25, respectively, and
tary environments can be used to test our under- assuming f = 4, Venus, Earth, and Mars have
standing of the factors that contribute to planetary equilibrium temperatures of 231 K, 255 K, and
habitability. The HZ is usually defined for surface 210 K, while their mean surface temperatures are
conditions only. Chemoautotrophic life, whose 737 K, 288 K, and 215 K.
metabolism does not depend on the stellar light, The orbital semimajor axis in the middle of the
can still exist outside the HZ, thriving in the habitable zone for a circular orbit, aHZ (in AU), is
interior of the planet where liquid water is derived by scaling the Earth-Sun system using
1050 Habitable Zone
Lstar =Lsun ¼ ðRstar =Rsun Þ2 ðTstar =Tsun Þ4 , so aHZ ¼ effective temperature of the star, the limits of
1 AUðLstar =LSun Þ0:5 and finally the habitable zone cannot be simply scaled to
the stellar luminosity; see ▶ Habitability,
Effects of Stellar Irradiation.
aHZ ¼ 1 AU ððL=Lsun Þ=Seff Þ0:5 : (2) Note that a planet found in the HZ is not
necessary habitable. Many factors may prevent
This formula (Kasting et al. 1993) assumes that surface habitability: lack of water or ingredients
the planet has a similar ▶ albedo to Earth, that it necessary for the emergence of life, gravity that is
rotates or redistributes the insolation as on Earth too small to retain a dense atmosphere against
(▶ Habitable Zone, Effect of Tidal Locking; gravitational escape (▶ Atmosphere Escape),
▶ Habitability, Effect of Eccentricity), and that and a lack of an active geology replenishing the
it has a similar greenhouse effect. Climate models atmosphere in carbon dioxide. The planet could
(▶ Atmosphere, Model 1D; ▶ GCM) are required have accreted a massive H2-He envelope that
to calculate the inner and outer limits of the HZ would prevent water from being liquid by keep-
for a planet with a given atmosphere. A revision ing the surface pressure too high, or it could have
of the calculations by Kasting et al. (1993) migrated to its current position (see, e.g., Valen-
reported values of Seff = 1.04 for the inner edge cia et al. 2006). In addition, the HZ assumes a
of the Sun HZ (runaway greenhouse limit) terrestrial planet with a water content large
(▶ Habitability, Effects of Stellar Irradiation) enough so that the surface can host liquid water
and 0.35 for the outer edge of the Sun HZ for any surface temperature between the temper-
(assuming a maximum greenhouse effect in the ature of the triple point of water, 273 K, and the
planet’s atmosphere; see Kopparapu et al. 2012, critical temperature of water, TC = 647 K
2013 and ▶ Spectral Type). (Kasting et al. 1993).
We assume here that the planet is rocky and
is dominated by a global carbonate-silicate
cycle that stabilizes the surface temperature Key Research Findings
and the CO2 level, like on Earth (Walker
1977). This implies that the planet is geologi- The limits of the circumstellar habitable zone
cally active and continuously outgasses CO2 (HZ) around a main sequence star are given in
and that carbonates form in the presence of Table 1. However, the limits of the HZ are known
surface liquid water, which may require conti- qualitatively, more than quantitatively. This
nental weathering. Without this stabilization, uncertainty is mainly due to the complex role of
the Earth would not be habitable over geologi- ▶ clouds and three-dimensional climatic effects
cal times (▶ Biomarkers Atmospheric, Evolu- not yet included in the modeling (▶ Atmosphere,
tion Over Geological Time). This in turn Model 1D; Atmosphere (▶ GCM)). Thus, planets
affects the composition of the atmosphere at slightly outside the computed HZ could still be
the outer edge of the habitable zone. As the habitable, while planets at habitable orbital dis-
amount of CO2 increases, the greenhouse effect tance may not be habitable because of their size
increases, warming the surface and releasing or chemical composition.
more CO2 from carbonates and from that A recent 3D study (LeConte et al. 2013)
dissolved in oceans. The feedback only works suggested that the inner HZ edge could be closer
up to the temperature where the increase in to the star (than predicted by some 1D models)
the albedo due to increasing CO2 concentration due to a Hadley cell response such that IR cooling
(producing more ▶ Rayleigh scattering and to space is shifted to lower altitudes where it is
▶ cloud formation; see, e.g., Forget and more efficient.
Pierrehumbert 1997) is stronger than the green- Given that stellar luminosity evolves during
house effect of CO2. Because of the relation the star’s lifetime, the concept of a continuously
between the albedo of the planet and the habitable zone (CHZ) has been introduced to
Habitable Zone 1051
Habitable Zone, Table 1 Properties and habitable zone limits of the main sequence stars of spectral type F-M
considered for the search of Earth analogs, assuming no other limits than runaway and maximum greenhouse for a rocky
planet with CO2 atmosphere – this model does not take increasing cloud fractions into account
Habitable zone (AU)b
a a a
Spectral type Effective temp. (K) Luminosity (L) Mass (M) Inner limitc Outer limitd
F 6,100–7,200 2.0–6.5 1.4–1.6 1.4–2.3 2.3–3.9
G 5,300–6,030 0.66–1.1 0.9–1.05 0.8–1.0 1.4–1.7
K 4,900–5,250 0.1–0.42 0.67–0.79 0.3–0.7 0.6–1.2
M 2,600–3,850 0.0012–0.077 0.06–0.51 0.04–0.3 0.1–0.6
a
Data from Ostlie and Carroll 1996
b
Data from the online HZ calculator at: http://depts.washington.edu/naivpl/content/hz-calculator
c
Runaway greenhouse limit calculated for planets around stars with Teff = 6,100 and 7,200 K for F stars, Teff = 5,300
and 6,030 K for G stars, Teff = 4,900 and 5,250 K for K stars, and Teff = 3,850 and 2,600 K for M stars
d
Maximum greenhouse limit for planets around stars with Teff = 6,100 and 7,200 K for F stars, Teff = 5,300 and
6,030 K for G stars, Teff = 4,900 and 5,250 K for K stars, and Teff = 3,850 and 2,600 K for M stars
mean the zone that remains habitable around a These regions have not been discounted as hab- H
star during a given period of time (Hart 1979). itable, but are extremely difficult to detect
remotely, even for moons in our own solar sys-
tem, and so will not be targeted by first-
Applications generation extrasolar planet characterization
missions. Only global atmospheric signatures
The concepts of the HZ and CHZ help define an are detectable remotely.
astronomical search zone for habitable planets
with remotely detectable species in their atmo-
sphere that indicate habitable conditions, such as Future Directions
surface liquid water. On a habitable planet where
the carbonate-silicate cycle is at work, the level The habitable zone concept is still evolving as we
of CO2 in the atmosphere depends on the amount learn more about planetary formation and evolu-
of greenhouse warming required to maintain tion and as we continue to improve the radiative
habitability and so depends on the orbital dis- transfer, ▶ clouds, and three-dimensional atmo-
tance: CO2 is a trace gas close to the inner edge spheric models (▶ Atmosphere-Ocean General
of the HZ but a major compound in the outer part Circulation Model) that allow more accurate cal-
of the HZ (Forget and Pierrehumbert 1997). culations of a planet’s temperature profile
Earth-like planets close to the inner edge are (▶ Atmosphere, Structure).
expected to have a water-rich atmosphere and
those at the outer edge of the habitable zone a
carbon dioxide-rich one. This is one of the first See Also
theories we can test with a first-generation space
mission. ▶ Albedo
There may be other habitable environments ▶ Atmosphere Escape
further from the star than the classical radiative ▶ Atmosphere, Model 1D
habitable zone, in what is known as the ▶ Atmosphere, Structure
“cryo-ecosphere” (see, e.g., Peña-Cabrera and ▶ Biomarkers Atmospheric, Evolution Over
Durand-Manterola 2004). This region could Geological Time
include environments made habitable by the ▶ Biomarkers, Spectral
delivery of tidal energy as would be the case ▶ Clouds
for a subsurface ocean on a giant planet’s moon ▶ Europa
(e.g., ▶ Europa or Titan-like environments). ▶ GCM
1052 Habitable Zone Around Binary Star Systems
▶ Habitability, Effect of Eccentricity Habitable Zone Around Binary Star
▶ Habitability, Effects of Stellar Irradiation Systems
▶ Habitable Zone, Effect of Tidal Locking Nader Haghighipour
▶ Main Sequence, Star Institute for Astronomy, University of Hawaii-
▶ Planetary Migration Manoa, Honolulu, Hawaii, HI, USA
▶ Rayleigh Scattering
▶ Spectral Type
Keywords
References and Further Reading Exoplanets; Habitability; Life
Forget P, Pierrehumbert H (1997) Warming early Mars
with carbon dioxide. Clouds that scatter infrared radi-
ation. Science 278:1273–1274 Definition
Hart MH (1978) The evolution of the atmosphere of the
Earth. Icarus 33:23–39 In a binary star system, a planet may in principle
Hart MH (1979) Habitable zones about main sequence
stars. Icarus 37:351–357
exist in an orbit around either of the stars or in an
Huang SS (1959) Occurrence of life outside the Solar orbit that surrounds both stars of the binary. In the
system. Am Sci 47:397–402 latter case, we refer to the planet as a
Huang SS (1960) Life outside the solar system. Sci Am ▶ circumbinary planet. The corresponding habit-
202(4):55
Kasting JF, Whitmire DP, Reynolds RT (1993) Habitable
able zone (HZ) may be assumed to be an annulus
zones around main sequence stars. Icarus 101:108–128 around the binary where an Earth-like planet
Kopparapu RK, Ramirez R, Kasting JF, Eymet V, Robin- (with similar atmospheric composition to that of
son TD, Mahadevan S, Terrien RC, Domagal- Earth) with a sufficient amount of water can per-
Goldman S, Meadows V, Deshpande R (2012) Habit-
able zones around main-sequence stars: new estimates.
manently maintain liquid water on its solid
Astrophys J 765(2), 131 surface.
Kopparapu RK, Ramirez R, Kasting JF, Eymet V, Robin-
son TD, Mahadevan S, Terrien RC, Domagal-
Goldman S, Meadows V, Deshpande R (2013) Erra-
tum: “Habitable Zones around Main-sequence Stars:
Basic Methodology
New Estimates”. Astrophys J 770, 82
Le Conte J, Forget F, Charnay B, Wordsworth R, Pottier To calculate the boundaries of the habitable zone
A (2013) Increased insolation threshold for runaway (HZ) around a binary star system (circumbinary
greenhouse processes on Earth-like planets. Nature
504:268–271
orbits), it is assumed that the HZ is an annulus
Ostlie DA, Carroll BW (1996) An introduction to modern around the binary where an Earth-like planet
stellar astrophysics. Addison-Wesley, USA (with similar atmospheric composition as that
Peña-Cabrera GVY, Durand-Manterola HJ (2004) Possi- of Earth) and sufficient amount of water can
ble biotic distribution in our galaxy. Adv Space Res
33:114–117
permanently maintain liquid water on its solid
Rasool SI, DeBergh C (1970) The runaway greenhouse surface. The locations of the boundaries of the
and the accumulation of CO: in the Venus atmosphere. HZ, and therefore the capability of the planet in
Nature 226:1037–1039 maintaining conditions for habitability, depend
Segura A, Kaltenegger L (2010) Search for habitable
planets. In: Basiuk VA, Navarro-González R (eds)
on the total flux received at the top of its atmo-
Astrobiology: emergence, search and detection of life. sphere. Since the atmosphere converts stellar
American Scientific Publishers, New York, pp 341–358 insolation to temperature structure and surface
Valencia D, O’Connell RJ, Sasselov DD (2006) Internal temperature of a planet, its interaction with the
structure of massive terrestrial planets. Icarus
181:545–554
stellar radiation plays an important role in con-
Walker JCG (1977) Evolution of the atmosphere. Mac- sidering whether a planet can maintain liquid
millan/Collier Macmillan, New York water on its surface and allow for detectable
Habitable Zone Around Binary Star Systems 1053
Habitable Zone Around Binary Star Systems, circles depict the boundary of orbital stability and the
Fig. 1 (a–d) Snapshots of the radial variations of the orbits of the currently known planets of the system
narrow (dark green) and empirical (light green) HZs (Figure from Haghighipour and Kaltenegger 2013)
around the Kepler 16 binary system. The red and blue
habitable conditions. This interaction strongly star’s effective temperature and f represents the
depends on the stellar spectral energy distribu- atmosphere’s cloud fraction. The contribution of
tion (SED), implying that stars with different each star of the binary to the total flux at the
energy distributions will contribute differently top of the planet’s atmosphere can be therefore
to the absolute incident flux at the top of the written as
planet’s atmosphere. To determine the contribu-
tion of a star taking into account its SED, it W i ðf, T i ÞFi ðf, T i Þ:
is necessary to define a spectral weight factor
Wi(f, Ti), where i = (Pr, Sec) denoting the pri- Here Fi ¼ Li =r 2Pli denotes the stellar flux and
mary and secondary stars, respectively, Ti is the Li is the luminosity of each star of the binary.
1054 Habitable Zone in Binary Stars Systems
The total flux received by the planet is therefore boundaries of the HZ. In this case, the determin-
given by ing factor is the averaged flux received by the
planet in one orbit around the binary. During this
LPr ðT Pr Þ LSec ðT Sec Þ time, the buffering effect of clouds should com-
W Pr ðf, T Pr Þ 2
þ W Sec ðf, T Sec Þ 2 ;
r Pl-Pr r Pl-Sec pensate for the effects of the temporary displace-
ments of the HZ and allow the planet to maintain
where rPlPr and rPlSec are the distances between a surface temperature conducive to liquid water.
the planet and the primary and secondary stars, Figure 1 shows snapshots of the circumbinary HZ
respectively. Defining HZ as a region where the of Kepler 16 system.
total flux received by an Earth-like planet at the
top of its atmosphere is equal to that of Earth
received from the Sun, the boundaries of References and Further Reading
circumbinary HZ can be calculated from
Haghighipour N, Kaltenegger L (2013) Astrophys J
777:166
LPr ðT Pr Þ LSec ðT Sec Þ Kaltenegger L, Haghighipour N (2013) Astrophys J
W Pr ðf, T Pr Þ 2
þ W Sec ðf, T Sec Þ 2 777:165
lx-Bin r Pl-Sec
Mueller T, Haghighipour N (2014) Astrophys J 782:26,
LSun article id
¼ :
l2x-Sun
where x-inner, outer boundary of the Hz.

In deriving this equation, it is assumed that the Habitable Zone in Binary Stars
primary star is at the center of the coordinates Systems
system (see the entry ▶ Circumbinary Planet) and
the boundaries of the HZ of the binary (lxBin) are Nader Haghighipour
measured with respect to this star. Institute for Astronomy, University of
The functional form of the spectral weight fac- Hawaii-Manoa, Honolulu, Hawaii, HI, USA
tor Wi(f, Ti) varies according to the models of the
Sun’s HZ. Haghighipour and Kaltenegger (2013)
have developed a detailed methodology for calcu- Keywords
lating this quantity. Mueller and Haghighipour
(2014) have utilized that methodology and created Exoplanets; Habitability; Life
a website with a user interface where one can input
the physical and orbital parameters of the binary,
and the website generates a snapshot of the Definition
circumbinary HZ corresponding to those orbital
elements. This website can be found at http:// In a binary star system, a planet may in principle
astro.twam.info/hz/. exist either in an orbit around a star or in an orbit
As implied by the above equation for calculat- that surrounds both stars in the system; in the
ing the HZ, the inner and outer boundaries of the former case, we refer to it as a ▶ circumprimary
HZ vary as the stars of the binary rotate around planet. To calculate the boundaries of the habit-
their center of mass. A gallery of the HZs of all able zone (HZ) around a star of a binary star
currently known circumbinary planets with system (circumprimary orbits), it is assumed
movies of their time variations can be found at that the HZ is an annulus around the star where
http://astro.twam.info/hz-ptype/. an Earth-like planet (with similar atmospheric
It is important to note that in reality, the hab- composition to that of Earth) and a sufficient
itability of an Earth-like planet in a circumbinary amount of water can permanently maintain liquid
HZ is independent of the fluctuations of the water on its solid surface.
Habitable Zone in Binary Stars Systems 1055
Habitable Zone in Binary 0.6

Stars Systems, M2M2 Binary
0.4 a = 1.5 (AU) , e = 0
Fig. 1 Boundaries of the
Distance [AU]
narrow (dark green) and 0.2
empirical (light green) HZs 0.0
in an M2-M2 (top), F0-F0 M M
(middle top), F8-M1 −0.2
(middle bottom), and −0.4
M1-F8 (bottom) S-type
−0.6
binary star systems. Note −0.5 0.0 0.5 1.0 1.5 2.0
that the primary is the star
Distance [AU]
at (0,0) (Figure from
Kaltenegger and 6
Haghighipour (2013)) F0F0 Binary
4 a = 15 (AU) , e = 0
Distance [AU]
2
0
F F
−2
−4
H
−6
−5 0 5 10 15 20
Distance [AU]
4 M1F8 Binary
a = 2.9 (AU) , e = 0
2
Distance [AU]
0
F
−2
−4
−8 −6 −4 −2 0 2 4 6 8
Distance [AU]
F8M1 Binary
4
a = 7.1 (AU) , e = 0
2
Distance [AU]
0
F
−2
−4
−8 −6 −4 −2 0 2 4 6 8
Distance [AU]
1056 Habitable Zone in Binary Stars Systems
6 M1F8 Binary 6
a = 4.24
4 e = 0.3 4
Distance [AU]
Distance [AU]
2 2
0 0
−2 −2
−4 −4
−6 −6
−5 0 5 −5 0 5
Distance [AU] Distance [AU]
6 6
4 4
Distance [AU]
Distance [AU]
2 2
0 0
−2 −2
−4 −4
−6 −6
−5 0 5 −5 0 5
Habitable Zone in Binary Stars Systems, of the F8 star on the extent of the HZ around the M1
Fig. 2 Boundaries of the narrow (dark green) and empir- primary while orbiting the primary starting from the top
ical (light green) HZs in an M1-F8 binary. Note that the left panel when the secondary is at the binary periastron
primary is the M1 star at (0,0). The panels show the effect (Figure from Kaltenegger and Haghighipour (2013))
Basic Methodology incident flux at the top of the planet’s atmosphere.

To determine the contribution of a star taking into
The locations of the boundaries of the HZ, and account its SED, it is necessary to define a spectral
therefore the capability of the planet in weight factor Wi(f, Ti), where i = (Pr, Sec)
maintaining conditions for habitability, depend denoting the primary and secondary stars, respec-
on the total flux received at the top of its atmo- tively, Ti is the star’s effective temperature, and
sphere. Since the atmosphere converts stellar inso- f represents the atmosphere’s cloud fraction. The
lation to temperature structure and surface contribution of each star of the binary to the total
temperature of a planet, its interaction with the flux at the top of the planet’s atmosphere can be
stellar radiation plays an important role in consid- therefore written as W i ðf , T i Þ. Here
ering whether a planet can maintain liquid water
on its surface and allow for detectable habitable Fi ¼ Li =r 2Pli
conditions. This interaction strongly depends on
the stellar spectral energy distribution (SED), denotes the stellar flux and Li is the luminosity of
implying that stars with different energy distribu- each star of the binary. The total flux received by
tions will contribute differently to the absolute the planet is therefore given by
Habitable Zone in Multi-star Systems 1057
LPr ðT Pr Þ LSec ðT Sec Þ liquid water. Figures 1 and 2 show snapshots of

W Pr ðf , T Pr Þ þ W Sec ðf , T Sec Þ 2 ; the HZs around stars of an FF, MM, and FM
r 2PlPr r PlSec
binary system.
where rPlPr and rPlSec are the distances
between the planet and the primary and second-
ary stars, respectively. Defining HZ as a region References and Further Reading
where the total flux received by an Earth-like
Haghighipouir N, Kaltenegger L (2013) ApJ 777:166
planet at the top of its atmosphere is equal to
Kaltenegger L, Haghighipour N (2013) ApJ 777:165
that of Earth received from the Sun, the bound- Mueller T, Haghighipour N (2014) ApJ 782, article id. 26
aries of circumprimary HZ can be calculated
from
LPr ðT Pr Þ LSec ðT Sec Þ

W Pr ðf , T Pr Þ 2
þ W Sec ðf , T Sec Þ 2 Habitable Zone in Multi-star Systems
lxBin r PlSec
LSun Nader Haghighipour H
¼ :
l2xSun Institute for Astronomy, University of Hawaii-
In deriving this equation, it is assumed that the
primary star is at the center of the coordinates
system (see the entry ▶ Circumprimary Planet) Definition
and the boundaries of the HZ of the binary
(lxBin) are measured with respect to this star. In a multiple star system, a planet may in princi-
The functional form of the spectral weight ple exist in an orbit around one or more stars.
factor Wi( f, Ti) varies according to the models Similarly, the habitable zone (HZ) of the system
of the Sun’s HZ. Kaltenegger and Haghighipour may be an annulus around one or more than one
(2013) have developed a detailed methodology of its stars, where an Earth-like planet (with sim-
for calculating this quantity. Mueller and ilar atmospheric composition to that of Earth)
Haghighipour (2014) have utilized that method- with a sufficient amount of water can perma-
ology and created a website with a user interface nently maintain liquid water on its solid surface.
where one can input the physical and orbital
parameters of the binary, and the website gener-
ates a snapshot of the circumprimary HZ Overview
corresponding to those orbital elements. This
website can be found at http://astro.twam.info/hz/ To calculate the boundaries of the habitable zone
As implied by the above equation for calculat- (HZ) in a multiple star system, it is assumed that
ing the HZ, the inner and outer boundaries of the the HZ is a region where an Earth-like planet
HZ vary as the stars of the binary rotate around (with similar atmospheric composition as that of
their center of mass. It is important to note that in Earth) and a sufficient amount of water can per-
reality the habitability of an Earth-like planet in a manently maintain liquid water on its solid sur-
circumprimary HZ is independent of the fluctua- face. The locations of the boundaries of the HZ,
tions of the boundaries of the HZ. In this case, the and therefore the capability of the planet in
determining factor is the averaged flux received maintaining conditions for habitability, depend
by the planet in one orbit of the binary. During on the total flux received at the top of its atmo-
this time, the buffering effect of clouds should sphere. Since the atmosphere converts stellar
compensate for the effects of the temporary dis- insolation to surface temperature, the interaction
placements of the HZ and allow the planet to of planet atmosphere with the stellar radiation
maintain a surface temperature conducive to plays an important role in considering whether a
1058 Habitable Zone in Multi-star Systems
a 6 b 6
5 5
4 4
3 3
2 2
A
Distance [AU]
Distance [AU]
1 1
KK A
0 0
−1 −1 KK
−2 −2
−3 −3
−4 −4
−5 −5
−6 −6
−7 −6 −5 −4 −3 −2 −1 0 1 2 3 4 5 6 −7 −6 −5 −4 −3 −2 −1 0 1 2 3 4 5 6
c 6 d 6
5 5
4 4
3 3
A
2 2
Distance [AU]
Distance [AU]
1 1
A KK
0 0
−1 −1
−2 KK −2
−3 −3
−4 −4
−5 −5
−6 −6
−7 −6 −5 −4 −3 −2 −1 0 1 2 3 4 5 6 −7 −6 −5 −4 −3 −2 −1 0 1 2 3 4 5 6
Habitable Zone in Multi-star Systems, Fig. 1 Time with respect to the horizontal line passing through zero
evolution of the HZ of the A–KK triplet in the KIC on the vertical axis. The dashed circles correspond to the
4150611 system when the orbit of the A star has a semi- outer boundary of the stability of planetary orbits. An
major axis of 5.48 AU and an eccentricity of 0.412. Dark animation of the HZ of this system can be found at
green corresponds to conservative HZ and light green http://astro.twam.info/hz-multi (Mueller and
denotes extended HZ. From top to bottom, the panels Haghighipour 2014)
correspond to the A star being at 0, 19, 86, and 180
planet can maintain liquid water on its surface the star’s effective temperature, and f represents
and allow for detectable habitable conditions. the atmosphere’s cloud fraction. The contribution
This interaction strongly depends on the stellar of each star of the system to the total flux at the
spectral energy distribution (SED), implying that top of the planet’s atmosphere can be therefore
stars with different energy distributions will con- written as
tribute differently to the absolute incident flux at
the top of the planet’s atmosphere. To determine W i ðf , T i ÞFi ðf , T i Þ:
the contribution of a star taking into account its
SED, it is necessary to define a spectral weight Here Fi ¼ Li =d2i denotes the stellar flux where Li is
factor Wi(f, Ti), where i denotes the ith star, Ti is the luminosity of the ith star of the system and di is
Habitable Zone in Multi-star Systems 1059
Habitable Zone in Multi-star Systems, Fig. 2 Screenshot of the interactive website http://astro.twam.info/hz for
calculating the HZ of binary and multiple star systems (Mueller and Haghighipour 2014)
1060 Habitable Zone in Multi-star Systems
Habitable Zone in Multi-

star Systems, Fig. 3 HZ
of a system of three 0.25
solar mass M stars in a
figure-eight orbit. From the
top to bottom, the figures
show the evolution of the
HZ of the system for one
complete revolution around
its center of mass. The
dashed circles show the
boundaries of the regions of
planetary stability around
each star (~0.101 AU).
As shown here, Earth-like
planets can have stable
orbits in a small region in
the empirical HZ around
each star. An animation of
the HZ of this system can
be found at http://astro.
twam.info/hz-multi
(Mueller and Haghighipour
2014)
Habitable Zone, Effect of Tidal Locking 1061
the distance from star i to the planet. The total flux of mass. However, it is important to note that in
received by the planet is therefore given by reality the habitability of an Earth-like planet is
independent of the fluctuations of the boundaries
X
N
Li of the HZ. In this case, the determining factor is
Ftotal ¼ W i ðT i Þ : the averaged flux received by the planet in one
i¼1 d2i
orbit. During this time, the buffering effect
of clouds should compensate for the effects of
Defining HZ as a region where the total flux
the temporary displacements of the HZ and
received by an Earth-like planet at the top of its
allow the planet to maintain a surface tempera-
atmosphere is equal to that of Earth received from
ture conducive to liquid water. Figure 3 shows
the Sun, the boundaries of a multi-star HZ can be
the HZ of an interesting triple star system in the
calculated from
famous figure-eight orbit (Mueller and
Haghighipour 2014).
X
N
Li LSun
W i ðT i Þ ¼ 2 :
i¼1 d 2i lxSun
See Also H
where x = (In, Out), and lxSun denotes the
boundaries of the Sun’s HZ. Any point in the ▶ Habitable Zone
HZ of the system will satisfy the condition ▶ Habitable Zone Around Binary Star Systems
▶ Habitable Zone in Binary Stars Systems
▶ Planets in Binary Star Systems
LSun X
N
Li LSun
W i ðT i Þ 2 ;
l2InSun i¼1
2
di lOutSun
In deriving this equation, it is assumed that the Haghighipouir N, Kaltenegger L (2013) Calculating the
center of mass of the system is at the center of the habitable zone of binary star systems. II: P-type bina-
coordinate system (Mueller and Haghighipour ries. Astrophys J 777, 166
Kaltenegger L, Haghighipour N (2013) Calculating the
2014). Figure 1 shows snapshots of the HZ of
habitable zone of binary star systems. I: S-type bina-
the triple star system KIC 4150611 from Kepler ries. Astrophys J 777, 165
Input Catalog. Mueller T, Haghighipour N (2014) Calculating the habit-
The functional form of the spectral weight able zones of multiple star systems with a new inter-
active website. Astrophys J 782, 26
factor Wi(f, Ti) varies according to the models of
the Sun’s HZ. Haghighipour and Kaltenegger
(2013) have developed a detailed methodology
for calculating this quantity. Mueller and
Haghighipour (2014) have utilized that methodol- Habitable Zone, Effect of Tidal
ogy and created a website with a user interface Locking
where one can input the physical and orbital
parameters of the binary, and the website gener- Lisa Kaltenegger
ates a snapshot of the circumbinary HZ Cornell University, Ithaca, NY, USA
corresponding to those orbital elements. This
website can be found at http://astro.twam.info/hz/.
Figure 2 shows a screenshot of this website. Keywords
As implied by the above equation for calcu-
lating the HZ, the inner and outer boundaries of Biomarkers; Habitability; Habitable zone;
the HZ vary as the stars rotate around their center M stars; Resonance; Tidal locking
1062 Habitat
Definition ▶ Habitable Zone

▶ Habitability of the Solar System
Tidal locking is a state of dynamical equilibrium ▶ Spectral Type
between a planet’s spin and orbital angular
momentum. Tidally locked planets on circular
orbits may become synchronous rotators,
presenting the same solid-body hemisphere to Dole SH (1964) Habitable planets for man. Blaisdell Pub-
the star. lishing, New York
Haberle RM, McKay C, Tyler D, Reynolds R (1996) Can
synchronous rotating planets support an atmosphere?
In: Doyle LR (ed) Circumstellar habitable zones.
Overview Travis House, Menlo Park, pp 29–41
Joshi M (2003) Climate model studies of synchronously
Tidally locked planets on circular orbits may rotating planets. Astrobiology 3:415–427
become synchronous rotators, presenting the Joshi MM, Haberle RM, Reynolds RT (1997) Simulations
of the atmospheres of synchronously rotating terres-
same solid-body hemisphere to the star. If the trial planets orbiting M dwarfs: conditions for atmo-
planet’s orbit is eccentric, either due to a primor- spheric collapse and the implications for habitability.
dial condition or maintained by interactions with Icarus 129:450–465
other planets, then the rotation period will be Kasting JF, Whitmire DP, Reynolds RT (1993)
Habitable zones around main sequence stars. Icarus
less than the orbital period and the planet will 101:108–128
not synchronously rotate. Planets interior to the
tidal lock radius may also become trapped in
spin-orbit resonances, where they complete an
integer ratio of rotations on their axis to orbits
around their star. For example, Mercury is Habitat
within the Sun’s tidal lock radius in a 3:2 spin-
orbit resonance, completing three rotations on its Felipe Gomez
axis every two revolutions (orbits) around the Centro de Astrobiologı́a (CSIC/INTA), Instituto
Sun. For an M dwarf (less massive and cooler Nacional de Técnica Aeroespacial, Torrejón de
than the Sun), the ▶ Habitable Zone is so close to Ardoz, Madrid, Spain
the star that the planets are likely to become
tidally locked within a relatively short time
after they form (Dole 1964; Kasting Synonyms
et al. 1993), but several resonances are possible.
Only a planet in a 1:1 resonance receives star- Ecosystem
light on only one hemisphere. Detailed models
have shown that this does not prevent habitabil-
ity for planets with even modestly dense atmo- Definition
spheres (Haberle et al. 1996; Joshi et al. 1997;
Joshi 2003). Even a tidally locked ‘Earth’ could Habitat is the location in an ▶ environment
remain habitable, provided that atmospheric which can be colonized by a biological popula-
cycles transport sufficient heat from the dayside tion. In nature, organisms live in association
to the nightside. with other organisms in assemblages called
populations. The components and number of
individuals in a habitat are governed by the
See Also resources and conditions that exist in the habitat.
Habitats differ markedly in their characteristics,
▶ Biomarkers, Spectral and a habitat that is favorable for the develop-
▶ Habitable Planet, Characterization ment of one organism may actually be harmful
Hadean 1063
for another organism. From an astrobiological determined as marking the lower limit of the
point of view, habitat is deeply related to Hadean by the Precambrian Subcommission on
the habitability concept which refers to the Stratigraphy in 2012. The ages have been
conditions that can support life in an adverse rounded to ~4600 Ma and 4000 Ma for the com-
surrounding. mencement and termination of the Hadean,
respectively, as shown on the International
Chronostratigraphic Chart.
See Also
▶ Biotope Overview
▶ Environment
The term Hadean was coined in 1972 by Preston
Cloud to cover that period of geological time
prior to formation of the earliest known rocks. It
was derived from Greek mythology where
HAC “Hades” first referred to the God of the Under- H
world and later to the Underworld itself. In
▶ Hydrogenated Amorphous Carbon geological usage, it is commonly equated with
“Hell,” based on the perception that the Earth
was extremely hot and turbulent at that time.
“Hadean” was an informal term for many
years, partly because of the diversity of views
Hadean as to its precise duration and to more recent
work that questioned whether much of this
Simon Wilde time period on Earth was really “hell like”. Its
Department of Applied Geology, Curtin starting point was variously taken as the forma-
University of Technology, Perth, Australia tion of the planetary nebula, the onset of accre-
tion, or the time from when the Earth was
largely solid. Its termination was placed at 3.8/
Keywords 3.9 Ga by some workers, the age of the most
ancient rocks in Greenland, although more
Precambrian; Jack Hills (Yilgarn, Western Aus- recent usage placed it at 4.0 Ga, based on the
tralia); Zircon age of the world’s oldest gneisses at Acasta,
Canada. Identification of zircon crystals up to
4.4 Ga, particularly from the Jack Hills in West-
Synonyms ern Australia, has led to the view that the Earth
had cooled down sufficiently by ~4.3 Ga to
Early earth; Priscoan support oceans and stable continents, based on
the study of oxygen and lithium isotopes in
pristine zircon domains. This raised the ques-
Definition tion as to whether the term “Hadean” should be
restricted to the period of time before the for-
The earliest eon of geological time, extending mation of oceans and continents, when the sur-
from accretion of the Earth (4.567 Ga) to the face of the Earth was too hot and unstable to
formation of the earliest known rocks. The allow liquid water to form and rocks to survive.
oldest rocks currently identified are components Furthermore, some workers who did not support
of the Acasta gneisses in the Northwest Terri- the use of “Hadean” preferred to extend the
tories of Canada (4.03 Ga), and this date was Eoarchean back to cover this period of
1064 Hadean Mantle
geological time. Further complicating the issue International Commission on Stratigraphy: http://www.
was that another term was in the literature for stratigraphy.org/index.php/ics-chart-timescale
Valley JW, Peck WH, King EM, Wilde SA (2002) A cool
essentially the same period of time: “Priscoan”, early earth. Geology 30:351–354
introduced by Brian Harland in 1989, although Van Kranendonk MJ, Altermann W, Beard BL, Hoffman
few published articles have used this name. PF, Johnson CJ, Kasting JF, Melezhik VA, Nutman
However, in 2012, following extensive discus- AP, Papineau D, Pirajno F (2012) A chronostra-
tigraphic division of the Precambrian: possibilities
sion by the Precambrian Subcommission, and challenges. In: Gradstein FM, Ogg JG,
“Hadean” was officially ratified by the Interna- Schmitz MD, Ogg GJ (eds) The geologic time scale
tional Commission on Stratigraphy. It was 2012. Elsevier, Boston, pp 299–392
accepted that it commenced with the onset of
formation of the Earth/Solar System and that it
terminated with the oldest known rocks pre-
served on Earth.
Planetary-scale geological processes took Hadean Mantle
place during the Hadean eon, from the accretion
of the Earth to its differentiation into a metallic Maud Boyet
core and a silicate mantle. Continental crust Université Blaise Pascal, Clermont-Ferrand,
developed earlier than previously thought, based France
on geological information retained by the Jack
Hills zircons. Several authors speculate that plate
tectonics could also have started during the Keywords
Hadean, yet geological evidence for this is still
controversial. A detailed account of these pro- Isotope systematics; Early Earth; Magma ocean;
cesses is reported in Earth, Formation and Mantle dynamics; Moon-forming impact
Early Evolution.
Definition
See Also
The structure and the composition of the Hadean
▶ Acasta Gneiss mantle are difficult to constrain due to the sparse
▶ Canadian Precambrian Shield geological record from this period. Isotope sys-
▶ Early Archean tematics show that the silicate Earth experienced
▶ Earth, Formation and Early Evolution global differentiation events during its infancy.
▶ Earth’s Atmosphere, History of the Origins Strong chemical fractionations and an extensive
▶ Geological Timescale loss of volatile elements are related to several
▶ Jack Hills (Yilgarn Craton, Western Australia) magma ocean stages, the last one following the
▶ Late Veneer giant impact forming the Moon. Although during
▶ Zircon the Hadean the mantle must have been hotter and
the convection more vigorous, early chemical
heterogeneities have not been erased until the
Archean, questioning the regime of the Hadean
mantle convection.
Bleeker W (2004) Towards a ‘natural’ time scale for the
Precambrian: a proposal. Lethaia 37:219–222
Cloud P (1972) A working model of the primitive earth. Overview
Am J Sci 272:537–548
Harland WB, Armstrong RL, Cox AV, Craig LE, Smith
AG, Smith DG (1990) A geologic time scale 1989. The Hadean eon corresponds to the earliest
Cambridge University Press, Cambridge, pp 1–28 period of the Solar System Earth’s history
Hadean Mantle 1065
starting from the beginning of solar system accre- convection relative to its modern convection
tion (4.568 Ga, Bouvier and Wadhwa 2010) to the regime driven by plate tectonics.
oldest preserved rocks dated by U-Pb systematics
(4.03 Ga, Acasta Gneiss, Bowring and Williams
1999). Geological records from this period that See Also
have been preserved over Earth’s history are
scarce and mostly consist of detrital zircons ▶ Acasta Gneiss
(Wilde et al. 2001). Rock samples directly ▶ Accretion
derived from the mantle are more difficult to ▶ Bulk Silicate Earth
date because they do not contain zircon which is ▶ Earth, Formation and Early Evolution
the mineral that is the most robust to metamor- ▶ Giant Impact
phism and alteration. A crustal section of Hadean ▶ Mantle
mafic crust has been discovered in Northern Can- ▶ Nuvvuagittuq (Porpoise Cove) Greenstone Belt
ada (▶ Nuvvuagittuq Greenstone Belt) with the
oldest units dated at 4.1 and 4.3 Ga (O’Neil
et al. 2012). The 4.3 Ga age is still debated H
(Roth et al. 2013), but these findings suggest
that the lower limit of the Hadean eon may have
Allègre CJ, Staudacher T, Sarda P, Kurz M (1983) Con-
to be redefined. Indirect information on the chem- straints on evolution of Earth’s mantle from rare gas
ical composition of the mantle and the chronol- systematics. Nature 303:762–766
ogy of the silicate differentiation is provided by Blichert-Toft J, Albarède F, Rosing M, Frei R, Bridgwater
D (1999) The Nd and Hf isotopic evolution of the
radioactive systematics that record parent-
mantle through the Archean. Results from the Isua
daughter chemical fractionation. Noble gas sys- supracrustals, West Greenland, and from the Birimian
tematics trace the mantle degassing, and the terranes of West Africa. Geochim Cosmochim Acta
xenon isotope signature is consistent with exten- 63:3901–3914
Bouvier A, Wadhwa M (2010) The age of the solar system
sive loss of volatile elements within the first
redefined by the oldest Pb-Pb age of a meteoritic
100 Ma of the Earth’s history (Allègre inclusion. Nat Geosci 3:637–641
et al. 1983; Yokochi and Marty 2005; Bowring SA, Williams IS (1999) Priscoan (4.00–4.03 Ga)
Mukhopadhyay 2012). 142Nd anomalies measured orthogneisses from northwestern Canada. Contrib
Mineral Petrol 134:3–16
in Archean samples and the large Hf isotope var-
Boyet M, Blichert-Toft J, Rosing M, Storey M, Télouk P,
iations measured in Hadean zircons and Archean Albarède F (2003) 142Nd evidence for early Earth
rocks are undisputable evidences for early silicate differentiation. Earth Planet Sci Lett 214:427–442
differentiation events during the first 150 Ma of the Caro G, Bourdon B, Birck J-L, Moorbath S (2003)
146Sm-142Nd evidence from Isua metamorphosed
Earth’s history (Blichert-Toft et al. 1999; Boyet
sediments for early differentiation of the Earth’s man-
et al. 2003; Caro et al. 2003; Kemp et al. 2010). tle. Nature 423:428–432
These isotope signatures are explained by multiple Kemp AIS, Wilde SA, Hawkesworth CJ, Coath CD,
generations of magma oceans during the accretion. Nemchin A, Pidgeon RT, Vervoort JD, DuFrane SA
(2010) Hadean crustal evolution revisited: new con-
The last important event of melting would be a
straints from Pb-Hf isotope systematics of the Jack
direct consequence of the Moon-forming ▶ giant Hills zircons. Earth Planet Sci Lett 296:45–56
impact, but the preservation of 182W anomalies Mukhopadhyay S (2012) Early differentiation and volatile
suggests that this catastrophic event did not induce accretion recorded in deep-mantle neon and xenon.
Nature 486:101–104
complete melting and homogenization of the
O’Neil J, Carlson RW, Paquette J-L, Francis D (2012)
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Guitreau M, Blichert-Toft J (2013) Inherited 142Nd
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1066 Hadley Cells
Touboul M, Puchtel IS, Walker RJ (2012) 182W evidence

for long-term preservation of early mantle differentia- Haeckel’s Conception of Origins
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Wilde SA, Valley JW, Peck WH, Graham CM (2001) Evi- of Life
dence from detritical zircons for the existence of con-
tinental crust and oceans on the Earth 4.4 Gyr ago. Stéphane Tirard
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Yokochi R, Marty B (2005) Geochemical constraints on
mantle dynamics in the Hadean. Earth Planet Sci Lett des Techniques EA 1161, Faculté des Sciences et
238:17–30 des Techniques de Nantes, Nantes, France
History
Hadley Cells
Ernst Haeckel (1834–1919) was a German
Lisa Kaltenegger zoologist, great supporter of the evolution theory.
Cornell University, Ithaca, NY, USA He developed his own theory of evolution
founded on Darwinism and also on some
Lamarckian principles. His theoretical thought
Definition was closely connected to his monistic
philosophical view.
The Hadley cell on the Earth is a circulation Regarding the origin of life, he proposed a
pattern that dominates the tropical atmosphere, scenario of ▶ abiogenesis based on protoplasmic
with rising motion near the equator, a compensa- theory and claimed that evolution of matter could
tory sinking motion in the subtropics, poleward lead to a form of albuminoidal protoplasm. He
flow in the upper troposphere (7–20 km above the called Monera the primitive entities coming before
surface) and equatorward flow near the surface. cells.
The Hadley circulation cells, one on each hemi-
sphere, cover about half of the Earth’s surface
area. The Hadley cells carry heat and moisture See Also
from the tropics to the northern and southern
mid-latitudes. Hadley cells are also present on ▶ Abiogenesis
other terrestrial bodies like Mars, Titan, or ▶ Darwin’s Conception of the Origins of Life
Venus. On a slowly rotating planet like Venus, ▶ Huxley’s Conception on Origins of Life
the Hadley circulation can extend almost to the ▶ Protoplasmic Theory of Life
poles. Circulation cells like the Hadley cells
probably exist also on other rocky exoplanets
and influence their climate substantially.
Haldane’s Conception of Origins
of Life
See Also
Stéphane Tirard
▶ Atmosphere, Structure Centre François Viète d’Histoire des Sciences et
▶ Scale Height des Techniques EA 1161, Faculté des Sciences et

Keywords
Chamberlain J, Hunten D (1987) Theory of planetary
atmospheres. Academic, Orlando Prebiotic soup
Half-Life 1067
History capable of reproduction; in other words, they

were the first genes (p. 107). Then he claimed
John Burdon Sanderson Haldane (1892–1964) that “when the whole sea was a vast chemical
was one of the famous British biochemists laboratory,” the condition for the formation of
and geneticists of the first part of the twentieth oily films and then primitive cells must have
century. He notably participated to the reflection been favorable.
about evolution questions, and he was one of the Haldane’s proposal, with Oparin’s one,
main actors on the synthesis of that issues. revealed the evolution of ideas on the origin of
During his career, he spoke several times life at the beginning of the twentieth century.
about the problem of the origins of life; however, Haldane, according to current biology, used his
he never worked as specialist of the topic. He biochemist and geneticist view to describe the
published a very important text on this matter in steps of a chemical process producing the most
1929, in The Rationalist Annual. This text is often simple living beings.
presented as linked to the text of Aleksandr
Ivanovich Oparin (1894–1980) (the Oparin’s
paper on origins of life was firstly published in See Also H
Russian in 1924), and it is often said to be Oparin-
Haldane’s theory. However, these two papers ▶ Miller, Stanley
were independently published. ▶ Oparin’s Conception of Origins of Life
Haldane gave a complete scenario describing ▶ Urey’s Conception of Origins of Life
primitive conditions on earth and steps of chem-
ical evolution from mineral to organic molecules.
The main reactions occurred in the primitive sea,
sort of complex chemical solution, named hot
dilute soup by Haldane. Haldane JBS (1929) The origin of life, rationalist annual.
His text began with a historical narrative about In: On being the right size and other essays. Oxford
spontaneous generation and by an analysis of the University Press, Oxford, 1991
nature of the bacteriophage, which he considered
as “a step beyond the enzyme on the road of life,
but it is perhaps an exaggeration to call it fully
alive” (p. 106). Half-Life
He suggested a complete scenario beginning
with the primitive atmosphere contained little or Francis Albarède
no oxygen, but carbon dioxide, and claimed that Ecole Normale Supérieure de Lyon, Lyon,
ammonia could be formed by the action of water France
on nitride contained in the crust. Moreover, in
the lack of ozone, ultraviolet rays could be
chemically active. One of his most important Synonyms
suggestion is that “when ultraviolet light acts
on a mixture of water, carbon dioxide, and Period (half-life period)
ammonia, a vast variety of organic substances
are made, including sugars and apparently some
of the materials from which proteins are built.” Definition
He claimed that in the current nature, such a
mixture would be eaten by microorganisms, but The half-life T1/2 is the time required to halve the
in the primitive conditions, in this “hot dilute number of atoms of a particular radioactive
soup,” big molecules could be synthesized. For ▶ nuclide. It relates to the ▶ decay constant l
him, “the first living or half-living thing” was through T 1=2 ¼ ln 2=l 0:69=l.
1068 Half-Major Axis
See Also also on the solution of the determination of lon-

gitude at sea. In geophysics Halley published
▶ Decay Constant charts of the variation of the compass of
▶ Earth, Age of the north Atlantic ocean, giving the first
▶ Geochronology charts with lines of equal magnetic declination
▶ Radioactivity plotted.
Among all his scientific contribution there one
that worth mentioning, his commitment and
enthusiasm in the publication of one of the fun-
damental works of modern science, the
Half-Major Axis
Philosophiæ Naturalis Principia Mathematica
(1687), of his friend Isaac Newton (1643–1727).
▶ Semimajor Axis
See Also
Halley, Edmond
▶ Comet
Portugal
Halogen
History Henderson James (Jim) Cleaves II

Edmund Halley (1656–1742) was a multifaceted Institute of Technology, Meguro–ku, Tokyo,
English scientist, fellow of the Royal Society Japan
(1678) and Royal Astronomer (1720), which Institute for Advanced Study, Princeton,
made significant contributions on mathematics, NJ, USA
astronomy, navigation and geophysics. Halley’s Blue Marble Space Institute of Science,
name will remain in history largely due to his Washington, DC, USA
work of 1705 on the motion of comets. The Center for Chemical Evolution, Georgia Institute
similarity that he founded between the orbital of Technology, Atlanta, GA, USA
elements for the comets seen in 1531, 1607,
and 1682 led him to suggest that this comet,
that same comet that would one day bear his Definition
name, would return in December 1758. In fact
the comet was observed on 25 December 1758 A halogen is a chemical element from Group
(very slightly later than Halley expected). 17 (in the IUPAC convention) (formerly VII,
Halley’s work in astronomy is not restricted VIIA) of the periodic table, composed of fluo-
to the movement of comets. He made important rine, chlorine, bromine, iodine, and astatine.
works in stellar astronomy, having been the first The man-made element 117 is predicted to be
to accurately cataloging the south sky hemi- a halogen. The Swedish chemist Berzelius
sphere. And he was also the first to propose coined the term “halogen” from the Greek
using transits of Mercury and Venus to deter- háls, “salt,” and gen, meaning “come to
mine the Earth-Sun distance and therefore the be” – for an element that produces a salt with a
scale of the solar system. He was very interested metal. The halogens are the only periodic table
Halophile 1069
group that contains elements in all three familiar definition is that of Kushner and Kamekura
states of matter at standard temperature and (1988) who classify organisms depending on the
pressure. At room temperature and pressure, salt concentration needed for optimum growth.
fluorine and chlorine are gases, bromine is a Thus, non-halophiles grow best in media
liquid, and iodine and astatine are solids. The containing less than 0.2 M salts, while halophiles
halogens show several trends as the atomic grow best in media containing from 0.2 to 5.2 M
number increases, including decreasing electro- dissolved salts. Halophiles can be further divided
negativity and reactivity and increasing melting into slightly halophilic (optimum growth
and boiling points. between 0.2 and 0.5 M salt), moderately halo-
In their elemental form, the halogens exist as philic (0.5–2.5 M salt), and extremely halophilic
diatomic molecules, but these are relatively (above 2.5 M salt).
unstable. Due to their high reactivity and electron
affinity, halogens are usually found in nature as
molecular compounds or as ions. Halogen anions, Overview
known as halides (e.g., Cl, Br, and F), and
oxoanions such as iodate (IO 3 ) are commonly All three domains of life include halophilic H
found in many minerals and are major compo- microorganisms. Archaeal halophiles, all belong-
nents of seawater. Halogenated organic coming to the Euryarchaeota, can be found among
pounds are also produced by living organisms. the methanogens and the members of the order
The halogens all form binary compounds with Halobacteriales that only includes extreme halo-
hydrogen, e.g., HF, HCl, HBr, and HI, which philes. Within this order, there is a single family
are strong Brønsted-Lowry acids. (the Halobacteriaceae) that includes 28 genera,
some of them of alkaliphilic organisms. The most
halophilic organism as well as the ones most
frequently found in hypersaline environments is
Halophile included within this family. Bacterial phyla such
as Cyanobacteria, Proteobacteria, Firmicutes,
Josefa Anton Actinobacteria, Spirochaetes, and Bacteroidetes
Department of Physiology, Genetics and are also halophiles (Oren 2008). Finally, some
Microbiology, University of Alicante, Alicante, eukaryotic organisms – plants, fungi, ciliates,
Spain and flagellates – are halophilic. The unicellular
green alga Dunaliella salina is the most halo-
philic known eukaryote. Normally, every halo-
Keywords phile has a relatively narrow range of salt
concentration that allows growth. The
Compatible solute; Hypersaline environment proteobacterium Halomonas spp. is an exception,
being able to grow over a very wide range of salt
concentration.
Synonyms Halophiles are usually found in phylogenetic
trees that also include non-halophilic relatives,
Salt-loving organisms but three large phylogenetically coherent groups
comprise only halophiles. Such is the case of
the aerobic ▶ Archaea from the order
Definition Halobacteriales (also known as haloarchaea),
the anaerobic fermentative ▶ bacteria from the
Halophile is an organism that needs high salt order Halanaerobiales, and the Gammaproteo-
concentrations for growth. A widely used bacteria from the family Halomonadaceae.
1070 Halophile
A group of phylogenetically related halophiles Alkaliphilic, psychrotrophilic, and thermophilic

within the Bacteroidetes is recovered from halophiles have been described (for instance,
hypersaline environments all over the world. the thermophilic bacterium Halothermothrix
Halophilic microorganisms use two different orenii, the archaeon Halorubrum lacusprofundi,
strategies to cope with the osmotic stress and members of the archaeal genus
stemming from the high ionic concentration of Natronobacterium).
their environments. Archaea of the order Moderately halophilic bacteria inhabit
Halobacteriales, anaerobic Bacteria of the hypersaline habitats with salinities around 10 %.
order Haloanaerobiales, and some members of Aquatic environments with higher, near-
the Bacteroidetes (i.e., Salinibacter ruber) accu- saturation salt concentrations may be densely
mulate high concentrations of inorganic ions populated with extremely halophilic microorgan-
(mostly potassium) in the cytoplasm. These isms, and this leads to bright red, orange, or
three groups include the most extreme halo- purple colorations due to the pigments of
philes. The so-called “salt-in” strategy requires these organisms. In some multipond solar
an adaptation of the whole intracellular machin- salterns, microbial diversity is found to decrease
ery to work in highly saline environments. All at higher salinities. However, the number of
other halophilic organisms keep their osmotic microorganisms increases with salinity, reaching
balance by producing and/or accumulating very high densities of up to 108 cells/ml at near-
“compatible” low-molecular-weight organic saturation concentrations. Most of these cells
solutes (see “compatible solutes”) whose con- correspond to organisms of the Archaea domain
centration is regulated according to the salinity although in some environments Bacteria can
of the environment. account for up to 30 % of the total counts.
Habitats for halophilic microorganisms Hypersaline environments all over the world
include salt marshes, salt lakes that may be alka- contain Archaea such as the square
line (Mono Lake, Wadi Natrun), or near neutral haloarchaeon Haloquadratum walsbyi and
(e.g., the Dead Sea, Great Salt Lake, Tuz Lake, related populations, as well as Halorubrum rep-
Chaka Lake, and some cold hypersaline lakes in resentatives. In the Bacteria domain, the
the Antarctica), man-made salterns either coastal Bacteroidetes Salinibacter spp. and related
or inland, hypersaline deep sea masses, salted phylotypes are frequently found in hypersaline
food, animal hides, saline soils, dessert salt settings close to saturation, although in some
crusts, and subterranean brines. cases Proteobacteria constitute the bulk of
These environments have been classified extremely halophilic Bacteria detected in these
depending on whether they originated through systems. At near-saturation concentrations, both
the evaporation of seawater or not, thus cell number and diversity of ▶ Eukarya decrease
talassohaline (e.g., solar salterns) and dramatically, leaving only Dunaliella and some
athalassoaline (e.g., the Dead Sea). extremely halophilic fungi in low numbers. At
Hypersaline environments are extreme habi- salinities above 10–15 %, large organisms dis-
tats and very high salt is not the only condition appear, with the exception of the brine shrimp
restricting biodiversity in these systems. They Artemia salina and the larvae of the fly Ephydra.
can also have high pH or, depending on their Hypersaline environments harbor the highest
geographical location, high or low tempera- numbers of viruses reported so far for aquatic
tures. In addition, some hypersaline environ- systems.
ments have very low nutrient availability or Most dissimilatory prokaryotic metabolisms
high concentrations of toxic compounds such can function in hypersaline environments
as heavy metals. Accordingly, halophilic organ- although a few of them have never been
isms may display additional extremophilisms. detected at salt concentrations above 10 %
Halotolerance 1071
(e.g., proton-reducing acetogens, methanogenesis Oren A (2008) Microbial salt at high salt con-
from acetate, or autotrophic nitrite oxidations). For centrations: phylogenetic and metabolic diversity.
Saline Syst 4:2
a detailed discussion of the energetics of Ventosa A (2006) Unusual micro-organisms from unusual
halophilism, see Oren (1999). habitats: hypersaline environments. In: Logan NA,
In addition to their evolutive and ecological Lappin-Scott HM, Oyston PCF (eds) SGM symposium
interest, halophilic microorganisms have some 66: prokaryotic diversity-mechanisms and signifi-
cance. Cambridge University Press, Cambridge
biotechnological applications. Haloarchaea pro-
duce bacteriorhodopsin that can be used as
photoactive material in optical devices as well
as polyhydroxyalcanoates, exopolysaccharides, Halotolerance
and halocins. Moderately halophilic bacteria are
used for the commercial production of their com- Josefa Anton
patible solutes ectoines and hydroxyectoines, Department of Physiology, Genetics and
which find application in the cosmetic industry Microbiology, University of Alicante, Alicante,
or the biological treatment of saline industrial Spain
waste effluents. Halophilic eukaryotes such as H
Dunaliella are used as a natural source of beta-
carotene in cosmetics and some food while the Keywords
shrimp Artemia salina can be used in
aquaculture. Compatible solutes; Ionic stress; Salt
See Also Synonyms
▶ Alkaliphile Salt tolerance

▶ Archaea
▶ Bacteria
▶ Compatible Solute Definition
▶ Eukarya
▶ Euryarchaeota Halotolerance is tolerance to ionic stress or the
▶ Extreme Environment ability of an organism to grow at salt concentra-
▶ Extremophiles tions higher than those required for growth.
▶ Halotolerance Halotolerant organisms are able to survive at
▶ Osmolite high salt concentrations but do not require these
conditions for growth.

Overview
DasSarma S, Arora P (2006) Halophiles. Encyclopedia of
life sciences. Wiley, London
Kushner DJ, Kamekura M (1988) Physiology of halophilic
Halotolerance is a relative term that refers to the
eubacteria. In: Rodrı́guez-Valera F (ed) Halophilic ability to survive or thrive at salt concentrations
bacteria. CRC, Boca Raton higher than those necessary for growth. A micro-
Oren A (1999) Bioenergetic aspects of halophilism. organism is considered extremely halotolerant if
Microbiol Mol Biol Rev 63:334–348
its growth range extends above 2.5 M salt. A 0.9 %
Oren A (2006) Life at high salt concentrations. In:
Dworkin M (ed) The prokaryotes: a handbook on the NaCl would be considered an isotonic solution by
biology of bacteria. Springer, New York most nonmarine and non-halophilic organisms. In
1072 Handedness
nature, halotolerant organisms can be found in Oren A (2008) Microbial salt at high salt concentrations:
settings such as saline waters and soils that are phylogenetic and metabolic diversity. Saline Syst 4:2
Roberts MF (2005) Organic compatible solutes of
inhabited by autochthonous halophilic microbiota halotolerant and halophilic microorganisms. Saline
or even in association with animals. Such is the Syst 1:5
case of microbiota such as Staphylococcus species
that live on human skin. Because of their exposure
to the salts in sweat, they are very halotolerant and
they can also grow well at NaCl concentrations as Handedness
high as 15 %. However, they can also grow in the
absence of salt and are thus said to be halotolerant ▶ Chirality
and not halophile. This property is exploited in the
design of selective growth media. Halotolerant
organisms have developed metabolic processes, Haphazardness
such as the accumulation of compatible solutes
that allow them to compensate the osmotic pres- ▶ Chance and Randomness
sure and continue to live in hyperosmotic
environments.
Salt tolerance can vary depending on nutri-
tional or environmental factors such as pH, tem- Hapten
perature, and redox potential. Since high salt has
been traditionally used as a way of preserving Henderson James (Jim) Cleaves II
food, halotolerant microorganisms can be a Earth–Life Science Institute (ELSI), Tokyo
cause of spoilage of salt-preserved food. Institute of Technology, Meguro–ku, Tokyo, Japan
The three domains of life include many exam- Institute for Advanced Study, Princeton,
ples of halotolerant microorganisms such as the NJ, USA
▶ bacteria of the genera Alteromonas, Lactoba- Blue Marble Space Institute of Science,
cillus, Bacillus, Myxococcus, and Pediococcus, Washington, DC, USA
as well as many cyanobacteria, some Center for Chemical Evolution, Georgia Institute
methanogenic and thermophilic archaea, and of Technology, Atlanta, GA, USA
halotolerant fungi such as Debaromyces,
Hansenula, Cladosporium, and Saccharomyces.
Synonyms
See Also
Partial antigen
▶ Adaptation
▶ Bacteria
Definition
▶ Compatible Solute
▶ Halophile
A hapten is a small molecule that can elicit an
▶ Osmolite
immune response when attached to a larger car-
rier molecule such as a ▶ protein. Neither the
References and Further Reading larger molecule nor the hapten may elicit the
response by themselves.
DasSarma S, Arora P (2006) Halophiles. Encyclopedia of
life sciences. Wiley, London
Oren A (2006) Life at high salt concentrations. In:
See Also
Dworkin M (ed) The prokaryotes: a handbook on the
biology of bacteria. Springer, New York ▶ Antibody
HAT 1073
Definition
Hard Landing
HARPS is a high-resolution spectrograph
Catharine A. Conley designed to achieve long-term stability enabling
NASA Headquarters, Washington, DC, USA Doppler velocity measurements at the level of
1 m/s, in order to derive radial-velocity orbits
for stars orbited by planets as small as the Earth.
Definition It is presently the only instrument scheduled on
the 3.6-m telescope at the European Southern
The term “hard landing” describes the disposition Observatory on La Silla in Chile. The spectro-
of a spacecraft on another planetary body that is graph is fed by fibers and the optics are enclosed
not controlled and could result in fragmentation in a vacuum tank, to eliminate the effects of
of the hardware. A loss of control resulting in a changes in the index of refraction at the echelle
hard landing is called a “crash”! For ▶ planetary grating due to changes in the atmospheric pres-
protection purposes, a hard landing is assumed to sure. The temperature of the instrument is sta-
result in the release of ▶ encapsulated bioburden bilized at the level of 0.001 K, which is also H
as well as the spreading of the ▶ exposed surface critical for the velocity stability. The wave-
bioburden. length calibration is provided by thorium-
argon hollow cathode lamps, but experiments
are underway with the use of Fabry-Perot
See Also etalons and/or Laser Combs as calibration
sources. HARPS has demonstrated the potential
▶ Encapsulated Bioburden to reach a velocity precision of perhaps
▶ Exposed Surface Bioburden 20 cm/s. HARPS and ▶ HIRES on Keck 1 in
▶ Planetary Protection Hawaii are now the premier instruments for
determining the masses of small planets. The
data from these two telescopes have been used
to detect several potentially habitable planets
around stars GJ 667C and Gliese 581. A copy
Hard Snowball
of HARPS for use in the northern hemisphere is
under construction.
▶ Snowball Earth
See Also
HARPS
▶ Doppler Shift
David W. Latham1 and Nader Haghighipour2
1 ▶ HIRES
▶ Radial-Velocity Planets
Cambridge, MA, USA
2
Synonyms HAT
High Accuracy Radial-velocity Planet Searcher ▶ HATNet

1074 HATNet
HATNet Hayabusa Mission
David W. Latham Anny-Chantal Levasseur-Regourd

Harvard-Smithsonian Center for Astrophysics, UPMC University of Paris 6/LATMOS-IPSL,
Cambridge, MA, USA Paris, France
Synonyms Keywords
HAT; Hungarian-made Automatic Telescope Asteroid; Regolith; Sample return

Network
Synonyms
Definition
MUSES-C
The Hungarian-made Automated Telescope
Network (HATNet) is a wide-angle ground-
based photometric survey for transiting Definition
exoplanets operated by the Smithsonian Astro-
physical Observatory. Large areas of the sky are Hayabusa (“peregrine falcon” in Japanese) is a
monitored robotically using seven small tele- scientific space mission developed by JAXA to
scopes (11 cm in diameter) located at two differ- explore a near-Earth asteroid and to return aster-
ent sites in Arizona (five telescopes) and Hawaii oidal soil, while validating new engineering tech-
(two telescopes), looking for planets that transit nologies (propulsion, landing, and atmospheric
their host stars. HATNet has discovered reentry). Launched in May 2003, the Hayabusa-
54 transiting exoplanets, many of which are 1 rendezvous with asteroid Itokawa was in Sep-
larger than Jupiter. An important exception is tember 2005. After 2 months of close observa-
HAT-P-11b, which is a ▶ hot Neptune in the tions, it descended and attempted to collect
field of view of NASA’s ▶ Kepler mission. An surface grains. It departed in 2007 and cruised
international extension of the project to the back to Earth. The sample canister returned
southern hemisphere known as HATSouth safely in July 2010. JAXA has confirmed the
became operational in 2009, with six 18-cm presence of tiny dust particles sampled on
telescopes located at three collaborating sites Itokawa.
(two telescopes at each site) in Chile, Australia,
and Namibia. Since 2009, this survey has dis-
covered 10 Jupiter-sized and larger extrasolar Overview
planets.
The Hayabusa-1 spacecraft consisted of a core
(1 1.6 2 m) and two solar paddles (5.7 m
width), with a total mass of 530 kg, including
See Also its propellant (for chemical propulsion) and
xenon (for electrical propulsion). It carried
▶ Exoplanets, Discovery four scientific instruments, the AMICA imaging
▶ Kepler Mission camera, the NIRS near-infrared spectrometer,
▶ Transiting Planets the LiDAR laser ranging instrument, and the
▶ TrES XRS X-ray fluorescence spectrometer. In addi-
▶ WASP tion, the probe embarked three instruments for
Haze Particles 1075
navigation: an optical camera (ONC), an elec- lows. Its shape may suggest that it is a contact
tromagnetic sensor (FBS), and a laser emitter to binary. Such results are to be taken into account
control the attitude during descent (LRF). As an when considering the orbital deflection tech-
engineering spacecraft, Hayabusa-1 tested two niques that should and could be implemented if
new developments, xenon ion engines and an a similar asteroid was discovered on a collision
autonomous navigation system, which both course with the Earth.
performed successfully. Following the overall success of this
The mission was quite complex and faced pioneering mission, Hayabusa-2 was launched.
major issues. Hayabusa-1 was launched on December 3, 2014, to rendezvous with a primi-
May 9, 2003, from Kagoshima Space Center tive C-type asteroid (162173), 1993 JU3, with
(now called Uchinoura), with an MV-5 solid similar goals and instruments. A major added
propellant rocket. Soon after, it was hit by a contribution is a lander, provided by DLR and
large solar flare that damaged the solar cells, CNES, designed to monitor the potential mag-
reducing the electrical power and the efficiency netic field of the asteroid and to perform in situ
of the ion engines. After an Earth swing-by in a microscopic characterization of the composi-
May 2004, Hayabusa arrived close to the tion of the asteroid, at a grain scale. H
targeted asteroid (25143) Itokawa in
mid-September 2005 and performed its thor-
ough remote characterization. In November See Also
2005, it released the Minerva mini-lander,
which unfortunately escaped Itokawa’s gravita- ▶ Itokawa Asteroid
tional pull and was lost in space. Hayabusa then ▶ Near-Earth Objects
attempted twice to touch down on the surface of
the asteroid, after having deployed a sampling
horn, later sealed after having tentatively col-
lected asteroidal dust. Hayabusa-1’s return References and Further Reading
maneuver started in April 2007, for a 3-year
Abe A et al (2006a) Near infra-red spectral results of
cruise back to the Earth. The return capsule, asteroid Itokawa from the Hayabusa spacecraft. Sci-
released from the main spacecraft 3 h before ence 312:1334–1338
reentering the Earth atmosphere, landed safely Abe S et al (2006b) Mass and local topography measure-
ments of Itokawa by Hayabusa. Science 312:
near Woomera, Australia, on June 13, 2010. It
1344–1347
was transferred to the JAXA curation facility. Demura H et al (2006) Pole and global shape of 25143
Preliminary analyses have confirmed that most Itokawa. Science 312:1347–1349
of the collected particles, which have sizes Fujiwara A et al (2006) The rubble-pile asteroid Itokawa
as observed by Hayabusa. Science 312:1330–1334
below 10 mm, have come from Itokawa.
Okada HT et al (2006) X-ray fluorescence spectrometry
The scientific observations made while of asteroid Itokawa by Hayabusa. Science 312:
Hayabusa-1 was hovering above Itokawa have 1338–1341
established that it is a very small body (major Saito J et al (2006) Detailed images of asteroid 25143
Itokawa from Hayabusa. Science 312:1341–1344
axes 530 290 210 m, with a density of
Yano H et al (2006) Touchdown of the Hayabusa space-
(1,925
160) kg m3), indicative of a rather craft at the Muses sea of Itokawa. Science
high porosity of about 40 %. This may suggest 312:1350–1353
that it is built from loose-packed rocks, held
together by their gravity. Itokawa presents a
significant topographic diversity, with a variety
of rough terrains scattered with boulders, as well
as featureless central areas, which likely result Haze Particles
from fine dust particles forming a smooth rego-
lith that has accumulated in local gravitational ▶ Aerosols
1076 HC4H
Definition
HC4H
▶ Hydrogen cyanide polymers – i.e., heteroge-
▶ Diacetylene neous solids formed upon spontaneous polymer-
ization of HCN – are likely to have been among
the first macromolecules on prebiotic Earth.
HCl Overview
▶ Chlorine Hydrides in the Interstellar Medium Hydrogen cyanide (HCN) polymerizes spontane-
▶ Hydrogen Chloride ously in the neat liquid and in solution. Due to the
abundance of hydrogen cyanide in the Universe
(Debes et al. 2008), the potential role of hydrogen
cyanide polymers in astrochemistry and the ori-
gin of life has provoked much speculation. Gas
HCl+
phase HCN is found both in interstellar molecular
clouds and in comets in the solar system.
The ▶ oligomerization of HCN was first
observed by Proust (1806), and analyses of the
soluble fraction date back to the late 1800s and
early 1900s (Lange 1863; Wippermann 1874;
HCN Dimer Bedel 1923). Volker was the first to propose a
structure for the insoluble ▶ polymer in 1960, a
▶ Cyanomethanimine so-called “ladder” structure formed by repeats of
the HCN dimer in the trans configuration (Volker
1960). The cis variation was subsequently offered
by Umemoto et al. (1987). A more provocative
model, from the standpoint of the origins of life,
HCN Polymer was suggested by Matthews, based on the obser-
vation of a-amino acids in polymer hydrolyzates;
Irena Mamajanov1 and Judith Herzfeld2 polymerization was proposed to occur via the
1
School of Chemistry and Biochemistry, Georgia HCN trimer aminomalononitrile and result in a
Institute of Technology, Atlanta, GA, USA structure with a -NCC- backbone that would
2
Brandeis University, Waltham, MA, USA form ▶ polypeptide upon contact with water
(Matthews et al. 1977). However, Ferris has
argued that the HCN tetramer, diaminoma-
Keywords leonitrile (DAMN), must be the “direct precursor”
to HCN polymers, since dimers and trimers are not
Addition polymer; Aminomalononitrile; expected to accumulate (Ferris et al. 1981).
Diaminomaleonitrile; Hydrogen cyanide; Ladder The above structures have been found to be
polymer; Polyheterocycle; Polyimine; inconsistent with recently acquired 13C and
15
Protopeptide N solid state NMR spectra of polymers formed
in neat HCN. These data suggest that the poly-
mers are formed by simple monomer addition,
Synonyms first in head-to-tail fashion to form linear, conju-
gated chains, and then laterally to form saturated
Azulmin; Hydrogen cyanide polymer two-dimensional networks (Mamajanov and
HCN Polymer 1077
HCN Polymer,
H2N
Fig. 1 Proposed structures N N N N
H 2N N
for the HCN polymers and
related compounds NH2 NH2 NH2 H2N H2N
H2N
N
NH2 NH2 NH2
N N N N N
Volker Structure Umemoto Structure
NH NH NC CN
H H H H H
H2N C C N C C N C C N
H 2N NH2
C C C
Diaminomaleonitrile (DAMN)
N N N
Protopeptide Structure H H
N N
N N N N
H2C C C C
H
N N N N
H H
N N HN N N NH
HN N NH
Addition Polymer Fragment Structures
N NH2
N
H
N
NH2 H N
N
N
NH2
poly-(aminoimidazole)
Herzfeld 2009a). This interpretation of the NMR See Also

spectra finds support in other information about
the polymerization of neat HCN, including the ▶ Biopolymer
presence of free radicals (Budil et al. 2003) and ▶ Exopolymers
the fragmentation pattern in a ▶ GC/MS ▶ Formamide
chemolysis study (Minard et al. 1998) (Fig. 1). ▶ GC/MS
A different kind of HCN polymer has been ▶ Heterocycle
shown to form upon mild heating of DAMN. 13C ▶ Hydrogen Cyanide
and 15N solid state NMR spectra, as well as the ▶ Insoluble Organic Matter
optical properties and electrical conductivity of the ▶ Nitrile
product, are consistent with an HCN trimer repeat ▶ Oligomer
unit. However, rather than forming a protopeptide, ▶ Oligomerization
the subunits cyclize to form poly-[aminoimidazole] ▶ Polymer
(Mamajanov and Herzfeld 2009b). ▶ Polypeptide
Despite extensive effort, there is no scientific ▶ Prebiotic Chemistry
consensus concerning the potential role of the ▶ Radical
HCN polymer in prebiotic evolution. ▶ Tholins
1078 HCNO Isomers
References and Further Reading Synonyms
Bedel C (1923) Sur un polymere de l’acide cyanhydrique. Cyanic acid; Fulminic acid; Isocyanic acid;
C R Acad Sci 176:168–171
Isofulminic acid
Budil DE, Roebber JL, Liebman SA, Matthews CN
(2003) Multifrequency electron spin resonance
detection of solid-state organic free radicals in HCN
polymer and a Titan tholin. Astrobiology 3(2): Definition
323–329
Debes JH, Weinberger AJ, Schneider G (2008) Complex
organic materials in the circumstellar disk of HR 4796- HCNO isomers are molecules having the same
A. Astrophys J 673:L191–L194 empirical formula (HCNO) but a different con-
Ferris JP, Edelson EH, Auyeung JM, Joshi PC nectivity. These isomers cannot be transformed
(1981) Structural studies on HCN oligomers. J Mol
from one to another without breaking a bond.
Evol 17(2):69–77
Lange O (1863) Ueber eine neue Verbindung von der
Zusammen-setzung der Cyanwasserstoffsaure.
Berichte 6:99 History
Mamajanov I, Herzfeld J (2009a) HCN polymers charac-
terized by solid state NMR: chains and sheets formed
in the neat liquid. J Chem Phys 130(13):134503 About 1825, Liebig and Gay-Lussac found out
Mamajanov I, Herzfeld J (2009b) HCN polymers that silver fulminate had the same empirical for-
characterized by SSNMR: solid state reaction of crys- mula as a compound called silver cyanate (more
talline tetramer (diaminomaleonitrile). J Chem Phys
correctly referred to as silver isocyanate), which
130(13)
Matthews C, Nelson J, Varma P, Minard R (1977) Wöhler had analyzed about a year earlier. The
Deuterolysis of amino-acid precursors – evidence for two compounds were clearly different and thus
hydrogen-cyanide polymers as protein ancestors. Sci- led to the concept of isomerism.
ence 198(4317):622–625
Minard RD, Hatcher PG, Gourley RC, Matthews CNZ
(1998) Structural investigations of hydrogen cyanide
polymers: new insights using TMAH Overview
thermochemolysis/GC-MS. Orig Life Evol Biosph
28:461–473
To date, four molecules are known with the
Proust JL (1806) Contributions on Cyanide. Ann Chim
Phys 60(1):233 empirical formula HCNO. The lowest-energy
Umemoto K, Takahashi M, Yokata K (1987) Studies on form is isocyanic acid, H─N=C=O; cyanic
the structure of HCN oligomers. Orig Life Evol Biosph acid, H─O─CN, is somewhat higher in energy
17(3–4):283–293
(103.2 kJ/mol); fulminic acid, H─CN─O, is
Volker TH (1960) Polymeric hydrocyanic acid. Angew
Chem Int Ed 72(11):379–384 considerably higher (295.9 kJ/mol); and
Wippermann R (1874) Ueber Tricyanwasserstoff, eine der isofulminic acid, H─O─NC, is higher still
blausaure polymere verbindung. Berichte 7:767 (352.2 kJ/mol). Fulminic acid is a linear mole-
cule; the others are all planar with the heavy
atoms being nearly linear and the H atom bent
trans to the heavy atom chain. A review of their
HCNO Isomers structural and spectroscopic properties and their
energetics was given by Teles et al. (1989). More
Holger S. P. M€uller recently, high-level quantum chemical calcula-
I. Physikalisches Institut, Universität zu Köln, tions of the energies have been performed by
Köln, Germany Schuurman et al. (2004).
Single-crystal X-ray crystallography has
shown that solid salts considered to be cyanates
Keywords are generally much better described as isocya-
nates. Consequently, adding strong acids to
Isomer these salts releases isocyanic acid (HNCO),
HD 189733b 1079
without clear evidence of formation of cyanic acid Schuurman MS, Muir SR, Allen WD, Schaefer HF III
(HOCN). HNCO was prepared in its pure form in (2004) Toward subchemical accuracy in computa-
tional thermochemistry: focal point analysis of the
the nineteenth century. In contrast, it appears as if heat of formation of NCO and [H, N, C, O] isomers.
HOCN can only be generated in situ. Interestingly, J Chem Phys 120:11586–11599
even though fulminic acid is much higher in Teles JH, Maier G, Hess BA Jr, Schaad LJ,
energy than cyanic acid, it is possible to prepare Winnewisser M, Winnewisser BP (1989) The CHNO
isomers. Chem Ber 122:753–766
HCNO in its pure form. Again, isofulminic acid
has to be generated in situ.
Rotational spectra of all four isomers have
been measured. Not surprisingly, isofulminic
acid was the last detected (Mladenović HCO+
et al. 2009). While HNCO was detected in space
in 1972, HCNO and HOCN were only detected ▶ Formyl Cation
very recently (see Br€unken et al. 2010).
HNCO can trimerize to cyanuric acid, a
six-membered ring with alternating C and H
N atoms.
The anion NCO is neither well described as HCOOCH3
isocyanate with the negative charge at the N atom
nor as cyanate with the negative charge at the ▶ Methyl Formate
O atom. Instead, the charge is delocalized to
essentially equal amounts over both atoms.
Thus, the CO bond has a formal bond order of
1.5 while that of the CN bond is 2.5. An infrared HCP
feature at 4.62 mm, observed in interstellar ices,
coincides with the strong asymmetric stretching ▶ Phosphaethyne
mode of the NCO anion and has been attributed
to this species. Unfortunately, features in solid-
state spectra are not particularly specific. As a
consequence, the assignment may well be correct
but is by no means certain. HD 189733b
Of the four isomers, the three lower-energy
ones have been detected in the ▶ interstellar David W. Latham1 and Nader Haghighipour2
1
medium: HNCO, HCNO, and HOCN. Harvard-Smithsonian Center for Astrophysics,
Cambridge, MA, USA
2
See Also Manoa, Honolulu, Hawaii, HI, USA
Keywords
References and Further Reading Planetary atmosphere; Transiting planet
Br€unken S, Belloche A, Martı́n S, Verheyen L, Menten

KM (2010) Interstellar HOCN in the Galactic center Definition
region. Astron Astrophys 516:A109
Mladenović M, Lewerenz M, McCarthy MC, Thaddeus
P (2009) Isofulminic acid, HONC: ab initio theory and HD 189733b is a transiting ▶ Hot Jupiter that is
microwave spectroscopy. J Chem Phys 131:174308 especially well suited for studies of its
1080 HD 189733b
HD 189733b, Fig. 1 The

light curve of HD 189733
observed with the Infrared
Array camera on the Spitzer
space telescope at 8 mm.
More than half the orbit is
covered. The deep event at
phase 0.0 is the transit of
the planet in front of the
star. The shallower event at
phase 0.5 is the secondary
eclipse when the planet
passes behind the star and
its thermal emission is
blocked. The depth of the
secondary eclipse indicates
that 0.34 % of the total
radiation of the system at
8 mm arises from the planet.
This sets the planet’s
dayside temperature at
1,212 K. The emission
increases slowly with phase
as the planet moves around
in its orbit and more of the
dayside comes into view. (b)
Provides an expanded view
of the upper portion of (a)
atmosphere. Spectroscopic observations with proved to be especially well suited for follow-up
the Hubble Space Telescope and ▶ Spitzer studies of its atmosphere because it is in a tight
Space Telescope have revealed the presence of orbit around a nearby star that is somewhat cooler
molecules such as water vapor and methane than the Sun, thus producing a favorable contrast
(Fig. 1). between the emission from the planet compared
to the star (Fig. 2).
History
Overview
Early results from radial-velocity surveys
suggested that gas giant exoplanets were being The parent star HD 189733 is significantly
found more frequently around stars with higher cooler than the Sun, with an effective
concentrations of heavy elements in their atmo- temperature of 4,980 K, corresponding to an
spheres than the Sun (e.g., see Santos et al. 2001; early K ▶ spectral type. The distance to the star
Fischer and Valenti 2005). This motivated some is only 19.3 pc, which makes it quite bright, with
teams to search for planets around samples of an apparent visual magnitude of 7.67. The period
metal-rich stars using radial velocities and then of the circular orbit is only 2.2186 days long, and
to look for transits with follow-up photometry. thus the planet is strongly heated by the incident
One of these efforts, the ELODIE metallicity- radiation from the star. These factors all combine
biased search for transiting Hot Jupiters to make HD 189733b especially well suited for
(da Silva et al. 2006), announced the discovery follow-up observations of the thermal emission
of HD 189733b (Bouchy et al. 2005). This planet from the planet’s atmosphere. Deming
HD 189733b 1081
HD 189733b, Fig. 2 The observed spectrum (black tri- departures are interpreted as additional absorption fea-
angles) and two theoretical spectra of the predominantly tures due to the presence of one or more other species in
molecular hydrogen atmosphere, showing the effects of addition to water. When considering only water and meth-
small amounts of water (blue) and methane in combina- ane, the theoretical spectrum best fitting the data was
tion with water (orange). The measured spectrum exhibits determined by binning the model (shown as white crosses)
significant differences at 1.7–1.8 mm and at 2.15–2.4 mm to the spectral resolution of the observations (Swain
from what is expected due to water vapor alone. These et al. 2008)
et al. (2006) first reported the detection of sec- dioxide to the list of proposed molecular detec-
ondary eclipses, using the Spitzer Space Tele- tions (Swain et al. 2009).
scope at 16 mm to measure a drop of 0.55 % in
the total thermal emission when the planet disap-
pears behind the star. Subsequently, Knutson See Also
et al. (2007) monitored the emission at 8 mm
over more than half the orbit and showed that ▶ Exoplanets, Discovery
the dayside temperature is 1,212 K, and the night- ▶ Gas Giant Planet
side is only 239 K cooler. This implies efficient ▶ Hot Jupiters
heat transfer from the dayside to the nightside and ▶ Hubble Space Telescope
thus dramatic weather in the atmosphere of HD ▶ Radial-Velocity Planets
189733b. ▶ Spitzer Space Telescope
Spectroscopy of the wavelength dependence ▶ Transiting Planets
of the transit depth in the near infrared with
the NICMOS instrument on the Hubble Space
Telescope by Swain et al. (2008) gave dramatic
confirmation for the earlier preliminary Bouchy F et al (2005) Astron Astrophys 444:L15
reports of the detection of water vapor in the da Silva R et al (2006) Astron Astrophys 446:717
H2-dominated atmosphere of HD 189733b and Fischer DA, Valenti JA (2005) Astrophys J 622:1102
suggested in addition the detection of methane. Knutson HA et al (2008) Nature 447:183
Santos N et al (2001) Astron Astrophys 373:1019
Subsequent NICMOS spectroscopy of second- Swain M et al (2008) Nature 452:329
ary eclipses added carbon monoxide and carbon Swain M et al (2009) Astrophys J 690:114L
1082 HD 209458b
unseen companion transited the star came right

HD 209458b on schedule. Follow-up photometry at times
predicted for transits by the spectroscopic orbit
David W. Latham1 and Nader Haghighipour2 with period 3.5247 days revealed dips in the light
1
Harvard-Smithsonian Center for Astrophysics, curve consistent with a ▶ gas giant planet 1.32
Cambridge, MA, USA times the radius and 0.64 times the mass of Jupi-
2
Institute for Astronomy, University of ter. For the first time it was possible to show that
Hawaii-Manoa, Honolulu, Hawaii, HI, USA the bulk density of a Hot Jupiter was approxi-
mately as expected for a gas giant planet. This left
little doubt that the growing population of radial-
Keywords velocity planet candidates, in the early days of
detecting extrasolar planets, must in fact be
Transiting planet planets.
The discovery of HD 209458b transits is
another example of simultaneity in science. Dis-
Definition covery papers from two independent teams were
published simultaneously and back to back in the
HD 209458b was the first exoplanet observed to same issue of the Astrophysical Journal Letters
transit its host star. (Charbonneau et al. 2000; Henry et al. 2000). The
two teams based their transit time predictions on
radial velocities measured independently with
History ▶ HIRES on the Keck I telescope and Elodie on
the 1.9-m telescope at the Observatoire de Haute
Arguments based on simple geometry predict that Provence. The follow-up photometry was accom-
about one out of ten planets orbiting stars similar plished with small telescopes, in the case of
to the Sun should be seen to ▶ transit their parent Charbonneau et al. (2000) using a 4-in. lens and
star if the orbital period is only a few days. HD ▶ CCD camera located in the parking lot of the
209458 was roughly the tenth star to show a High Altitude Observatory in Boulder, Colorado.
radial-velocity orbit implying a possible Hot Subsequent observations of transits with the STIS
Jupiter companion, so the discovery that the instrument on the Hubble Space Telescope
HD 209458b, Fig. 1 The

light curve for the transit of 1.000
HD 209458b obtained with
the STIS instrument on the
Hubble Space Telescope.
Observations from several 0.995
relative flux
orbits were combined to

produce full phase
coverage of the transit light
curve (Brown et al. 2001) 0.990
0.985
–0.10 –0.05 0.00 0.05 0.10

time from center of transit (days)
Heat Flow, Planetary 1083
illustrated (Fig. 1) the spectacular quality of light References and Further Reading
curve that could be achieved with a large tele-
scope in space (Brown et al. 2001). Brown TM et al (2001) Hubble space telescope time-
series photometry of the transiting planet of HD
209458. Astrophys J 552:699
Charbonneau D et al (2000) Detection of planetary transits
Overview across a sun-like star. Astrophys J 529:45 L
Charbonneau D et al (2002) Detection of an extrasolar
planet atmosphere. Astrophys J 568:377
HD 209458 is a star very similar to the Sun in
Henry GW et al (2000) A transiting “51 Peg-like” planet.
almost all respects. Because it is relatively nearby, Astrophys J 529:41 L
only 47 pc from the Sun, it has the relatively bright Vidal-Madjar A et al (2003) An extended upper atmo-
apparent visual magnitude of 7.65. This makes it sphere around the extrasolar planet HD209458b.
Nature 422:143–146
especially attractive for follow-up observations,
such as spectroscopy of the atmosphere of the
planet HD 209458b, although the planetary candi-
date ▶ HD 189733b is somewhat better in this
regard because its host star is cooler. The same HD 44179 H
STIS spectra that yielded the spectacular light
curve when integrated over all the available wave- ▶ Red Rectangle
lengths (Brown et al. 2001) were subsequently
reanalyzed at much finer spectral resolution, show-
ing the presence of sodium in the planet’s atmo-
sphere (Charbonneau et al. 2002). This was the Heat Flow, Planetary
first detection of the atmosphere of an exoplanet.
Subsequently, hydrogen was detected by Vidal- Tilman Spohn
Madjar et al. (2003). Deutsches Zentrum f€ur Luft- und Raumfahrt
Although the bulk density of HD 209458b (DLR), Institut f€ur Planetenforschung, Berlin,
matches more or less the predictions of models Germany
for gas giant planets, the density is somewhat
lower than expected. Apparently the radius of the
planet is expanded compared to those predicted by Keywords
the models. For a while HD 209458b was anoma-
lous in this regard, but as more transiting Hot Radioactive heating; Heat transfer
Jupiters were discovered, additional examples of
inflated gas giants were identified. Although a
number of possible physical mechanisms for Synonyms
inflating the radii of these planets have been
suggested, the phenomenon remains puzzling. Geothermal flux; Heat flux
See Also Definition
▶ Exoplanets, Discovery The heat flow in the most general sense is the rate
▶ Gas Giant Planet of heat transfer per unit area across a given sur-
▶ HD 189733b face. The heat transfer mechanism could be radi-
▶ HIRES ation, convection, or conduction. It is measured
▶ Hot Jupiters in W m2. In geophysics and in planetary geo-
▶ Transit physics, the surface heat flow is more narrowly
▶ Transiting Planets defined as the heat conducted through the ▶ crust
1084 Heat Flow, Planetary
across the planetary surface. In the (older) geo- continental shields where heat flow is low. The
physical literature, the heat flow unit (hfu) is average value is 84 W m2, and the variation is
sometimes used, with 1 hfu = 106 cal between a few tens to a few hundreds mW m2.
cm2 s1 = 41.84 mW m2. Heat flow has been measured on the Moon at two
sites by the Apollo missions. The values are
12 and 21 mW m2 (Langseth et al. 1976; Warren
Overview and Rasmussen 1987). These values are still sub-
ject of debate. A recent review can be found in
Heat is generated in the interior of planets and Siegler and Smrekar (2014). No other planetary
satellites by the decay of radioactive elements, by heat flow values are available, but heat flow mea-
exothermic phase transitions and chemical reac- surements have been repeatedly proposed (e.g.,
tions, and by the dissipation of mechanical Spohn et al. 2001).
energy upon, for example, planetary contraction In the outer solar system, the intrinsic lumi-
or differentiation and tidal deformation. The rate nosities of ▶ Jupiter, ▶ Saturn, ▶ Uranus, and
of heat loss of a ▶ planet or satellite provides ▶ Neptune have been measured. These are
constraints on the amount of heat being produced 5.44
0.43, 2.01
0.14, 0.042
0.047 (but
in the planet and the temperature distribution in not smaller than 0), and 0.433
0.046 W m2,
the interior. The heat loss can – in principle – be respectively (e.g., Guillot and Gautier 2015 for a
measured from space with infrared radiometers review). Dissipation of gravitational energy
as the heat radiated by the planet. The latter released upon contraction is the major source of
quantity is also called the planet’s intrinsic heat in the interior of Jupiter. The heat flow value
▶ luminosity (similar to the luminosity of a for the other ▶ giant planets, in particular, the
▶ star). In the inner solar system, however, the difference in heat flow between Uranus and Nep-
luminosity is dominated by the solar heat that is tune, is not well understood. The intrinsic lumi-
absorbed in the atmosphere and in the surface nosity of the Jovian satellite ▶ Io has been
▶ rock and radiated back into space. (The contri- measured to be about 2 W m2 (Spencer
bution by the reflected sunlight can be separated et al. 2000). It is widely agreed that the heat is
in the electromagnetic spectrum.) Therefore, the due to tidal dissipation (compare Tides).
heat flow out of the interior of a ▶ terrestrial
planet must be measured at a depth where pertur-
bations due to the solar radiation and thermal See Also
perturbations in the atmosphere are sufficiently
small. This p depth
ffi is a few times the thermal
ffiffiffi ▶ Apollo Mission
kt
skin depth p with k the thermal diffusivity ▶ Crust
and t the period of the thermal perturbation. ▶ Differentiation, Planetary
The period t ranges from a planetary day ▶ Earth
(or ▶ sol) to centuries if climatic effects are ▶ Giant Planets
taken into account. Usually, a depth of a few ▶ Heat Transfer, Planetary
meters will be sufficient but depending on the ▶ Io
required accuracy. Measuring the heat flow then ▶ Jupiter
requires measuring both the thermal conductivity ▶ Luminosity
and the temperature gradient. Many heat flow ▶ Moon, the
measurements have been made on the ▶ Earth, ▶ Neptune
and heat flow maps have been compiled (e.g., ▶ Planet
Pollack et al. 1993; see also Jaupart and ▶ Rock
Mareschall 2015 for a review). The variation of ▶ Satellite or Moon
the heat flow reflects volcanic and orogenic prov- ▶ Saturn
inces and chains where heat flow is high and thick ▶ Sol
Heat Transfer, Planetary 1085
▶ Solar System, Inner microbiology, this technique is used to count the

▶ Star spores, generally in a liquid sample. The combi-
▶ Terrestrial Planet nation of high temperature with a short exposure
▶ Tides, Planetary time (i.e., 80 C for 10 min) kills the vegetative
▶ Uranus forms of the mesophilic ▶ bacteria, leaving
mesophilic and thermophilic spores alive. Within
such a short time, the microorganisms have not
References and Further Reading enough time to produce new spores. When
cooled, the solution is spread in or on a nutritive
Guillot T, Gautier D (2015) Giant planets. In: Spohn T, media to perform the count of the viable micro-
Schubert G (eds) Treatise on geophysics, vol 10. organisms left. The result of this standardized
Jaupart C, Mareschall JC (2015) Heat flow and thermal
▶ assay represents the spores which were present
structure of the lithosphere. In: Watts AB, Schubert G in the original sample.
(eds) Treatise on geophysics 6:217–253
Langseth MG, Keihm SJ, Peters K (1976) Revised lunar
heat flow values. P Lunar Planet Sci C 7:3143–3171 H
Pollack HN, Hurter SJ, Johnston SR (1993) Heat flow See Also
from the Earth’s interior. Rev Geophys 31:267–280
Siegler MA, Smrekar SE (2014) Lunar heat flow:
Regional prospective of the Apollo landung sites. ▶ Bacteria
J Geophys Res. doi:10.1002/2013JE004453 ▶ Endospore
Spencer JR, Rathburn JA, Travis LD, Tamppari LK, ▶ Microorganism
Barnard L, Martin TZ (2000) Io’s thermal emission ▶ Spore
from the Galileo photopolarimeter-radiometer.
Science 288:1198–1201
Spohn T, Ball A, Seiferlin K, Conzelmann V,
Hagermann A, Kömle NI, Kargl G (2001) A heat
flow and physical properties package for the surface
of Mercury. Planet Space Sci 49:1571–1577 Heat Transfer, Planetary
Warren PH, Rassmussen KL (1987) Megaregolith insula-
tion, internal temperatures and bulk uranium content
of the Moon. J Geophys Res 92:3453–3465
Tilman Spohn1 and Lisa Kaltenegger2
1
Germany
2
Heat Flux
▶ Heat Flow, Planetary Keywords
Atmospheres; Conduction; Convection;

Planetary interiors; Plate tectonics; Radiation;
Heat Shock Stagnant lid
Catharine A. Conley
NASA Headquarters, Washington, DC, USA Definition
Heat is transferred in planetary and satellite inte-

Definition riors and atmospheres from regions of high tem-
perature to regions of lower temperature. The heat
The term “heat shock” describes a fast and short transfer mechanisms are heat conduction, convec-
heat treatment. For ▶ planetary protection and tion, and radiation.
1086 Heat Transfer, Planetary
Overview t smaller than unity, the atmosphere is transpar-

ent and does not absorb efficiently (see Atmo-
In planetary interiors, heat is generated by the spheric Structure, Phase Change: Latent Heat,
decay of the ▶ radiogenic isotopes 235U, 238U, Thermal Inversion, Opacity).
237
Th, and 40K. In addition, latent heat may be Heat Conduction. In gases, heat is conducted
released and consumed through phase transitions. via the ▶ diffusion of molecules. Molecules with
The dissipation of kinetic energy of ▶ planetesi- high kinetic energy diffuse from regions of high
mals colliding with the protoplanets during plan- temperature to regions of lower temperature.
etary accretion has heated the planetary interiors, Interaction between molecules transfers energy
in some cases up to the melting temperatures of and thereby heat. In solids, energy is transferred
their outer layers. The formation of the cores in via lattice vibrations. In a perfect solid, vibra-
terrestrial planets through differentiation has fur- tional energy would travel with sound speed,
ther heated the deep interiors. Since planetary and heat would be rapidly dissipated. Imperfec-
surfaces are colder – their temperature being tions in the solid slow the process of energy
determined by the rate of solar radiation and the transfer, such that heat flows analogous to the
atmosphere ▶ greenhouse effect – heat is trans- diffusion of matter. One model of conductive
ferred to these surfaces from the deep interior. heat transfer associates vibrational energy with
The resulting flow of heat may be associated with particles called phonons (it can be shown that
the conversion of heat into mechanical work and vibrational energy must be quantized). The trans-
the conversion of heat into ▶ magnetic field fer of heat can then be modeled as diffusion in the
energy. Moreover, the cooling may result in plan- phonon gas similar to diffusion in a real gas.
etary contraction. The dominant heat transfer In planetary atmospheres, heat conduction is
mechanisms in planetary interiors are heat con- largely unimportant (except for the uppermost
duction and convection (for a discussion of heat layers) with convection and radiation being more
transfer in planetary interiors, see, e.g., Schubert efficient heat transfer mechanisms. In planetary
et al. 2001). interiors, heat is conducted in regions that are
In a typical substantial planetary atmosphere, immobile on planetary timescales. One example
heat is transferred by convection or radiation in layer is the stagnant ▶ lithosphere of planets such
the lower atmosphere (more specifically, the as ▶ Mars. On ▶ Earth where the lithosphere is
▶ troposphere and ▶ stratosphere) and heat con- broken up in plates that move laterally, vertical
duction and radiation in the upper atmosphere heat transfer is still conductive, while lateral heat
(more specifically, the mesosphere and thermo- transfer is convective. Other prominent layers of
sphere). The pressure level at the radiative- conductive heat transfer are the boundary layers of
convective boundary (the tropopause) in the mantle convection (compare Fig. 1 below and
lower atmosphere depends on the composition Schubert et al. 2001).
of the atmosphere and the ▶ opacity of the Mathematically,
!
the (conductive) flow of
major atmospheric constituents. Above this heat F can be described by
level (i.e., at lower pressures), the outgoing
!
energy is transported by radiation until heat con- F ¼ k∇T (1)
duction becomes important. The radiative-
convective boundary occurs when the atmo- where ! T is temperature and k is thermal conduc-
sphere becomes optically thin to thermal radia- tivity. F is a vector quantity having a value and a
tion. Usually, this translates into a column optical direction.
depth, t, approximately equal to unity. For t The thermal conductivity can be both a func-
approximately equal to unity or larger, the atmo- tion of pressure and temperature.
sphere is opaque, and the radiative energy is Radiation. Radiation is a heat transfer mech-
reabsorbed by the surrounding media, hence con- anism in regions that are transparent to electro-
vection is the dominant form of heat transfer. For magnetic waves and is particularly important in
Heat Transfer, Planetary, Fig. 1 Color-coded temper- stagnant layer, heat is transferred by conduction. In
ature distributions in stagnant lid (a) and mobile lid con- mobile lid convection, the cold near-surface layer partic-
vection (b) for strongly temperature-dependent viscosity ipates in the convection. Arrows indicate the direction of
after numerical calculations by Stein and Hansen (2008). flow. Mobile lid convection is a mode of convection H
Blue indicates low temperatures and red high tempera- largely analogous to plate tectonics (Courtesy of C. Stein
tures. In stagnant lid convection, the flow is confined to and U. Hansen)
occur underneath a cold near-surface stagnant layer. In the
planetary atmospheres (see radiation in planetary mechanism of the transfer of heat generated in
atmospheres and Thomas and Stamnes 1999). For the interior of the planet to the same cold sur-
the highly absorbing planetary interiors, radiation face. It is the same mechanism of heat transfer
is usually integrated into the mechanism of heat from the interior of the hot core to the (colder)
conduction thereby increasing the thermal con- core-mantle boundary and from the warm sur-
ductivity at high temperatures. face of the planet to the colder regions in the
Convection. Thermal convection is a conse- atmosphere. In the atmosphere, convection
quence of thermal expansion with the density of includes large- and small-scale rising and sink-
gases and liquids decreasing with increasing tem- ing of air masses and smaller air parcels. These
perature. Consider a layer of fluid (or gas) vertical motions effectively distribute heat and
sandwiched between a hot plate at the bottom moisture throughout the atmospheric column.
and a cold plate at the top. Fluid near the hot The mechanism also applies in the interiors of
plate will absorb energy, and its temperature ▶ giant planets and in oceans. It is termed free
will rise. As a consequence of the density convection because the flow is not driven by
decreasing with temperature, the fluid will external forces. Forced convection in planets
become increasingly buoyant with respect to the may be driven by, e.g., ▶ tides, planetary and
colder fluid above and eventually will rise if the in the atmosphere by variations in solar
!
irradia-
buoyancy is large enough to overcome inertia and tion. The convective flow of heat F across a
internal friction. As it rises, it will transport heat surface in a fluid is given by
that it may exchange with the fluid through which
it rises and – in particular – with the fluid near the ! !
cold top plate. The heat transfer is thus from the F ¼ ∇ u rcp T (2)
hot to the cold plate. In a similar thought exper-
!
iment, heat can be generated within the fluid, where u is the velocity vector, r is the local
cause buoyancy, and again be transported to the density, and cP the heat capacity at constant
cold plate. The above is – in principle – the mech- pressure.
anism of heat transfer from the hot core through Heat transfer by free convection can be char-
the mantle to the cold planetary surface and the acterized by a set of dimensionless parameters.
1088 Heat Transfer, Planetary
The first and most important is the Rayleigh b

Ra
number Ra. For fluids heated from below, it is Nu ¼ (5)
Racrit
defined as
where Racrit is the critical Rayleigh number for
gaDTd 3 the onset of convection and b is a constant, the
Ra (3)
nk value of which is between 0.2 and 0.33 depending
on boundary conditions and the temperature and
In addition to already defined quantities, g is the pressure dependence of material properties. Sim-
acceleration due to gravity, a is the thermal expan- ilarly, for internally heated convection,
sion coefficient, DT is the temperature difference
between the top and bottom boundary, d is the g
layer thickness, n is the kinematic ▶ viscosity, kT Ra
y ¼ (6)
and k(rcp) is the thermal diffusivity. Qd 2 Racrit
For internally heated fluids, DT is replaced
with Qd 2/k, where Q is the heat production rate where T is the internal temperature and g is
per unit volume. The term g aDT in the nominator between 0.2 and 0.25.
of Eq. 3 measures the thermal buoyancy. Note that The parameters a, n, and k are pressure and
there are two diffusivities in the denominator. The temperature dependent. Of particular importance
thermal diffusivity k measures the loss of buoy- for planetary mantles – as numerical calculations
ancy due to thermal conduction, and the kinematic and laboratory experiments have shown – is the
viscosity n – the momentum diffusivity – measures pressure and temperature dependence of the vis-
the deceleration of the fluid due to momentum cosity (see ▶ Rheology, Planetary Interior for a
diffusion. discussion). Note that the solid mantles of terres-
The steady-state heat transfer rate by convec- trial planets and satellites can be regarded as
tion through a fluid heated from below is mea- extremely viscous fluids for processes operating
sured by the Nusselt number Nu on geological timescales (i.e., on timescales of
millions of years or more). The extremely large
qd viscosity in the mantle causes inertia forces (such
Nu (4) as the Coriolis force) to be negligible and the flow
kDT
to be laminar, although at very large Rayleigh
with q the heat flow across the top surface. The numbers, the flow tends to become increasingly
Nusselt number is the ratio between the surface chaotic (see, e.g., Ricard 2007). Within a ▶ plan-
heat flow q and the heat flow that would be etary interior, and in particular through the litho-
transferred by conduction kDT/d; since convec- sphere, the outer boundary layer of the planet, the
tion is an instability that sets in when the heat can viscosity may vary over many orders of magni-
be more efficiently transferred by convection, tude. A strongly temperature-dependent viscosity
Nu 1. In a fluid heated from within, the surface can result in the formation of a stagnant lid
heat flow in the steady state equals Qd, and the (compare Fig. 1). The pressure dependence can
Nusselt number as defined above is always unity. be speculated to lead to a second stagnant layer in
The efficiency of convective heat transfer is then the deep interior of the planet. The formation of a
measured by the value of the internal tempera- mobile lid requires the variation of the viscosity
ture: the lower the temperature at which the con- to be sufficiently small. The formation of a plate-
vection is operating, the more efficient is the like mobile lid (the fluid dynamical representa-
internally heated convection. tion of ▶ plate tectonics) may require a
Experiments with fluids and numerical solu- viscoelastic rheology in which the yield strength
tions of the field equations of fluid mechanics may matter (compare Fig. 2). But it is not
show that there is a power law relationship completely understood how plate tectonics on
between the Nu and the Ra numbers the Earth works and what the exact conditions
Heat Transfer, Planetary, Yield stress

Fig. 2 Qualitative phase
Episodic
diagram for convection of a
fluid with viscoelastic Stagnant lid
rheology including strongly
temperature-dependent Stagnant lid
viscosity. The figure is
based on numerical
calculations by K. Stemmer
and colleagues. It
qualitatively shows how the
stability fields of the
stagnant lid and mobile lid
Episodic
modes of convection Mobile lid
depend on the Rayleigh
number, the viscosity
contrast across the fluid,
and the yield strength for Viscosity contrast
Mobile lid
the transition from elastic to

plastic deformation. There
H
is a transitional regime
termed episodic in which a
stagnant lid episodically
becomes mobile (Courtesy
of K. Stemmer) Rayleigh number
(Planetary radius)
for the stability of plate tectonics are. The latter ▶ Giant Planets
makes it, of course, difficult to predict plate tec- ▶ Greenhouse Effect
tonics on other planets. ▶ Interior Structure, Planetary
While the viscosity and its dependence on ▶ Jupiter
pressure and temperature is thought to be a most ▶ Magnetic Field
important parameter for mantle convection, vis- ▶ Mars
cosity is of little importance for convection in the ▶ Opacity
planetary atmosphere and the core and in the ▶ Planetesimals
interiors of the giant planets. In these, inertia ▶ Planet Formation
forces, in particular the Coriolis force, matter ▶ Plate Tectonics
and the flow is mostly turbulent. The Coriolis ▶ Radiogenic Isotopes
force provides helicity (corkscrew-like motion) ▶ Rheology, Planetary Interior
for the flow in the core which is a prerequisite for ▶ Satellite or Moon
dynamo action. In the atmosphere, the Coriolis ▶ Stagnant Lid Convection
force causes the vortices that dominate the ▶ Stratosphere
weather system on, e.g., the Earth. In the giant ▶ Terrestrial Planet
planets, the Coriolis force is again taken to be the ▶ Tides, Planetary
reason for the observed vortices (see ▶ Jupiter). ▶ Troposphere
See Also References and Further Reading
▶ Core, Planetary Pierrehumbert RT (2010) Principles of planetary climate.

▶ Diffusion Ricard Y (2007) Physics of mantle convection. In:
▶ Dynamo, Planetary Bercovici D, Schubert G (eds) Treatise on geophysics,
▶ Earth vol 7. Elsevier, Amsterdam, pp 31–88
1090 Heat-Stable DNA Polymerase
Schubert G, Turcotte D, Olson P (2001) Mantle convec-

tion in the Earth and planets. Cambridge University Heavy Hydrogen
Press, Cambridge, p 940
Seager S (2010) Exoplanet atmospheres: physical pro-
cesses. Princeton University Press, Princeton ▶ Deuterium
Stein CA, Hansen U (2008) Plate motions and the viscos-
ity structure of the mantle – insights from numerical
modeling. Earth Planet Sci Lett 272:29–40
Thomas GE, Stamnes K (1999) Radiative transfer in the
atmosphere and ocean. Cambridge University Press, Heavy Ion
Cambridge
▶ HZE Particle
Heat-Stable DNA Polymerase

Heavy Nucleus
▶ Taq Polymerase
▶ HZE Particle
Heavy Charged Particle
▶ HZE Particle Heavy Primary
▶ HZE Particle
Heavy Element
Daniel Rouan 6 Hebe

Meudon, France Philipp Gast
Asteroids and Comets, Institute of Planetary
Research, German Aerospace Center, Berlin,
Definition Germany
In the astrophysical context, a heavy element is

any element heavier than helium. The term Synonyms
“metal” is also used by astronomers. The defini-
tion stems from the fact that only helium and 1947 JB
hydrogen (and small quantities of lithium and
beryllium) were produced in the few first minutes
after the big bang: all other elements were pro- Definition
duced afterward, through stellar processing
(nuclear reactions or supernovae). 6 Hebe is a large asteroid residing in the inner
main belt with a semimajor axis of approximately
2.425 AU. Its orbit is inclined by about 14.748
See Also degree and has an eccentricity of 0.201. With
these orbital parameters, it is situated near the
▶ Metallicity 3:1 resonance with Jupiter and the v6 secular
▶ Nuclear Reaction resonance. This proximity makes it likely for
▶ Primordial Nucleosynthesis ejecta of Hebe, which can be caused by impacts
Hematite 1091
of smaller asteroids, to come into near-earth

regions by mediation of Jupiter or Saturn. Hebe
is therefore suggested to be one of the main
contributors for the terrestrial meteoroid flux of
H chondrites and the IIE iron meteorites.
With its diameter of 185.18 km and a mass
between 1.2 and 1.4 1019 kg, it has a similar
average density as Mars even though it is much
smaller. Hebe has been classified to be of spectral
class S(IV), of which it is a relatively atypical Hematite, Fig. 1 Mosaic of photos taken by the Mars
member because of its low spectral slope and UV rover Opportunity showing hematite “blueberries” spher-
color. ules spread over the Martian soil grains (Photo credit:
NASA)
History
Definition H
Hebe was discovered in 1847 by Karl Ludwig
Hencke in Driesen (Germany) as the sixth aster- Hematite is an iron oxide of chemical formula
oid ever to be found. Fe2O3 (trigonal crystal system). Hematite forms
in a large variety of geological environments and
is mined as one of the main ores of ▶ iron on
See Also Earth. Hematite is the major iron oxide in the
▶ banded iron formations (BIFs), which were
▶ Asteroid Belt, Main deposited during global oxidation events. It is
▶ Half-Major Axis also found in high-grade ore bodies in ▶ meta-
▶ Meteorites morphic rocks due to contact ▶ metasomatism
▶ Orbital Resonance and occasionally as a sublimate on lavas. Hema-
tite can be associated to liquid water, particularly
in hot spring deposits. For this reason, hematite
on planetary surfaces is a potential indicator of
the past presence of water. Hematite spherules,
Helium Nuclei
called “blueberries” (Fig. 1) due to their blue hue
in false-color images released by NASA, were
▶ Alpha Rays
found by the ▶ Mars exploration rover Opportu-
nity at Meridiani Planum. Embedded in a sulfate
salt matrix or loose on the surface (Fig. 1), those
hematite spherules might derive from accretion
Hematite underwater, though alternative formation pro-
cesses such as meteoric impact or volcanic pro-
Daniele L. Pinti cesses have also been advocated.
Montréal, QC, Canada See Also

Synonyms ▶ Goethite
▶ Great Oxygenation Event
Kidney ore ▶ Hydrothermal Environments
1092 Heme
▶ Iron photosynthesis. Melvin Calvin suggested that

▶ Iron Cycle heme may have served as a primitive peroxidase
▶ Iron Oxides, Hydroxides and Oxy-hydroxides prior to the formation of proteins.
▶ Jarosite
▶ Mars
▶ Metamorphic Rock See Also
▶ Metasomatism
▶ Oxygenation of the Earth’s Atmosphere ▶ Iron
▶ Photosynthesis
▶ Porphyrin
▶ Protein
Heme

Earth–Life Science Institute (ELSI), Tokyo HEPA Filters
Japan Catharine A. Conley
Institute for Advanced Study, Princeton, NJ, NASA Headquarters, Washington, DC, USA
USA
Washington, DC, USA Synonyms
of Technology, Atlanta, GA, USA High-efficiency particulate absorbing filters;
High-efficiency particulate air filters; High-
efficiency particulate arresting filters
Definition
Definition
A heme is a ▶ protein prosthetic group that con-
sists of an ▶ iron ion chelated in the center of a
These filters are manufactured and tested to
▶ porphyrin. A substantial fraction of porphyrin-
ensure that filtered air has undergone a reduction
containing metalloproteins have heme as their
in the level of particulate contamination. An
prosthetic group. Heme-containing proteins
HEPA filter, i.e., H14, is certified and tested to
have a number of biological functions including
filter 99.975 % of the particles with a minimal
diatomic gas transport (e.g., hemoglobin), chem-
size of 0.3 mm. Such filters are used in space
ical catalysis (e.g., peroxidase), and electron
industry to isolate some volumes from a dusty
transfer (e.g., cytochrome). The heme iron serves
or contaminated environment. For ▶ planetary
as a source or sink of electrons during redox
protection they are used in ▶ clean rooms to filter
chemistry; however, in several peroxidases, the
the incoming air and in BSL four laboratories to
porphyrin molecule also serves as an electron
filter both air inlets and outlets.
source. In the transportation of diatomic gases,
the gas binds to the heme iron, often inducing a
conformational change in the surrounding
protein. See Also
The original function of heme-containing pro-
teins may have been electron transfer in primitive ▶ Biological Safety Level
sulfur-based ▶ photosynthesis pathways in ances- ▶ Clean Room
tral cyanobacteria before the evolution of oxygenic ▶ Planetary Protection
Herschel Mission 1093
science payloads. In November 1993, the ESA

Herschel Mission Science Programme Committee (SPC) decided
that FIRST would be implemented as the fourth
Göran L. Pilbratt “Cornerstone” (CS4) mission. The science pay-
ESA/ESTEC/SRE-S, European Space Agency load was selected in 1997–1999, the major
(ESA), Science Support Office, Noordwijk, industrial contractors in 2000–2001, and in
The Netherlands April 2001, the industrial activities commenced.
Meanwhile, in December 2000, FIRST was
renamed Herschel in recognition of the 200th
Keywords anniversary of the discovery of infrared light
by William Herschel. The science instruments
Astrochemistry; Galaxy formation; Galaxy evo- were delivered and integrated in 2007; final
lution; Infrared galaxies; Infrared observatories; spacecraft integration, testing, and verification
Interstellar chemistry; Interstellar dust; Interstel- activities took place in ESA’s ESTEC Test Cen-
lar medium; Interstellar molecules; Molecular tre from January 2008 to January 2009, when the
clouds; Protostars; Space observatories; Star for- Herschel spacecraft was flown to Kourou, H
mation; Stellar evolution French Guiana, for the launch campaign. Her-
schel was launched on May 14, 2009, operated
from a large halo orbit around the second
Definition Lagrangian point (L2) in the Sun-Earth system,
and performed its final science observation on
The Herschel Space Observatory is a ▶ Euro- April 29, 2013, when it ran out of superfluid
pean Space Agency (ESA) facility for liquid helium coolant. The spacecraft was put
far-infrared and submillimeter astronomy, pro- in its heliocentric “parking” orbit, where it will
viding unprecedented observing capabilities for reman indefinitely, and finally turned off on June
the ~55–670 mm spectral range. Herschel was 17, 2013. Herschel observers will be supported
launched on May 14, 2009, and performed its in the exploitation of Herschel data throughout
final science observation on April 29, 2013, hav- the postoperation phase, which ends by the end
ing carried out ~23,500 h of successful science of 2017.
observing. All data and associated data products
are publicly available to the worldwide astron-
omy community through the Herschel Science Overview
Archive. The main science objectives address
▶ interstellar medium physics, star formation Mission
and evolved ▶ stars, and galaxy evolution. The Herschel Space Observatory is the fourth
“Cornerstone” mission in the European Space
Agency (ESA) “Horizon 2000” science plan.
History Herschel is a space facility for far-infrared
and submillimeter astronomy, providing
Herschel has a long history. Conceived of as the unprecedented observing capabilities in the
Far Infrared and Sub-millimetre Telescope ~55–670 mm spectral range. It offers a much
(FIRST) and proposed to ESA in November larger telescope and extends the spectral cover-
1982 in response to a call for mission proposals age to longer wavelengths compared to earlier
issued in July 1982, it was incorporated into the infrared missions such as the Infrared Astronom-
ESA “Horizon 2000” long-term plan in 1984. ical Satellite (IRAS), the Infrared Space Obser-
Over the years, it was the subject of multiple vatory (ISO), Akari, and Spitzer; additionally, it
studies adopting a variety of mission designs, bridges the wavelength gap to ground-based
spacecraft configurations, telescopes, and submillimeter facilities and provides an
1094 Herschel Mission
Herschel Mission, Fig. 1 Left: Herschel being prepared showing the three science instrument focal plane units
for acoustic testing in the ESTEC Test Centre in June on the optical bench on top of the helium tank. Also
2008, providing a good view of the telescope. Right: shown is the “Russian doll” structure minimizing “para-
A computer cut-away image of the payload module sitic” heat loads (ESA)
observational database for follow-up observa- Spacecraft and Telescope

tions, in particular with ALMA. The Herschel spacecraft (Fig. 1) provides the
Herschel was launched on May 14, 2009, by appropriate working environment for the science
an Ariane 5 ECA, together with Planck, from instruments, points the telescope with required
Europe’s spaceport in Kourou, French Guiana. accuracy, autonomously executes the observing
It was operated autonomously in a large timeline, and performs onboard data handling
quasihalo orbit around the second Lagrangian and communication with the ground. It has a
point (L2) in the Sun-Earth/Moon system, with modular design, consisting of the “payload mod-
a daily ground contact period of normally about ule” (PLM) supporting the telescope, the sun-
3 h. Herschel successfully observed until it ran shade/sunshield, and the “service module”
out of superfluid helium coolant on April (SVM). The mission lifetime is determined by
29, 2013, permanently terminating any further the cryostat lifetime, required to be 3.5 years.
taking of science data. The spacecraft was put in The PLM is dominated by the cryostat vacuum
its heliocentric “parking” orbit, where it will vessel (CVV) from which the superfluid liquid
reman indefinitely, and finally turned off on helium (SLHe) tank is suspended, surrounded by
June 17, 2013. Herschel observers will be three vapor-cooled shields to minimize parasitic
supported in the exploitation of Herschel data heat loads. The optical bench with the three
throughout the postoperation phase, which ends instrument focal plane units (FPUs) is supported
by the end of 2017. on top of the tank, containing ~335 kg SLHe at
Herschel Mission, Table 1 Herschel science instrument Basic Methodology

main characteristics
Principal Operations and Science Ground Segment
Acronym Instrument investigator The mission operations, conducted by the Mis-
PACS Imaging camera A. Poglitsch, sion Operations Centre (MOC), of Herschel are
and grating MPE, Garching (D) now completed. The science operations are con-
spectrometer,
spectral coverage tinuing and conducted by five centers:
55–205 mm
SPIRE Imaging camera M. Griffin, • The Mission Operations Centre (MOC), pro-
and FTS U. Cardiff (UK) vided by ESA
spectrometer, • The Herschel Science Centre (HSC), provided
spectral coverage
194–671 mm by ESA
HIFI Heterodyne F. Helmich, SRON, • Three dedicated Instrument Control Centres
spectrometer, Groningen (NL) (ICCs), one for each instrument, provided by
spectral coverage the respective PIs
157–212 and
• The NASA Herschel Science Center (NHSC), H
240–625 mm
provided by ▶ NASA
launch. A phase separator allows a continuous The HSC, supported by the ICCs and the
evaporation of the liquid into cold gas, keeping NHSC (primarily for the US science community),
the FPUs and their detectors at their required will continue to support the community to exploit
temperatures. The telescope is 3.5 m in diameter, the science data throughout the postoperation
was made of silicon carbide (SiC), had an inflight phase, ending by the end of 2017. Data
operating temperature of about 88 K, and had processing software and science archive will con-
very low emissivity. It provided a total wavefront tinue to be improved, refined products ingested
error (WFE) of less than 5 mm in the focal surface into the archive, and documentation enhanced,
interfacing to the instruments. finally yielding the Herschel Legacy Archive.
The SVM houses “warm” payload electronics
on four of its eight panels and provides the nec- Observing
essary “infrastructure” for the satellite such as Herschel was operated as an observatory, with a
power, altitude and orbit control, the onboard fraction of guaranteed time and open time avail-
data handling and command execution, commu- able to the worldwide community. Astronomers
nications, and safety monitoring. It also provides applied for open time on Herschel competitively
a thermally controlled environment, which is by responding to calls for proposals
critical for some of the instrument units. Finally, (Announcements of Opportunity – AOs), provid-
the SVM also provides mechanical support for ing the science objectives, the definition of the
the PLM, the sunshield/sunshade, and a thermal actual observations being proposed, and addi-
shield to thermally decouple the PLM from the tional information. All proposals were reviewed
SVM, and it ensures the main mechanical load by the independent Herschel Observing Time
path during the launch. Allocation Committee (HOTAC). The observa-
tions by the successful proposers are then carried
Science Instruments out. A small amount of the open time was allo-
Herschel has three science instruments: the Pho- cated as discretionary time.
todetector Array Camera and Spectrometer Given that Herschel would not have the bene-
(PACS), the Spectral and Photometric Imaging fit of an all sky survey for much of its wavelength
REceiver (SPIRE), and the Heterodyne Instru- coverage, it was decided early on that large
ment for the Far Infrared (HIFI). Their main so-called key programs (KPs) in the form of
characteristics are provided in Table 1. “large spatial and spectral surveys” should be
Herschel Mission, Fig. 2 ESA Herschel and NASA l = 10 and stretching ~1.5 above and below the
Spitzer view of ~1/30 of the Galactic Plane. This map plane (Observations from S. Molinari Herschel Hi-GAL
combines Spitzer/MIPS 24 mm (blue), Herschel/PACS Key Programme and S. Carey Spitzer MIPSGAL Pro-
160 mm (green), and Herschel/SPIRE 250 mm (red) data, gram. Image processing by H. Bouy (CAB CSIC),
it covers 12 3.3 deg^2 centred at galactic longitude E. Bertin (IAP), and B. Merı́n (ESA))
selected prelaunch and executed early in the mis- emission peaks in the 100–200 mm range. Gas in
sion. The AO for KPs was followed by two the ISM emits a profusion of spectral lines in the
inflight AOs. Herschel wavelength range.
Herschel successfully observed until it ran out Herschel particularly addresses the formation
of superfluid helium coolant. Since October and evolution of stars and galaxies and the phys-
29, 2013, all Herschel science observations are ics and chemistry with the ISM which they form
publicly available from the Herschel Science from and interact with. In addition, Herschel has
Archive (HSA). The HSA offers over 37,000 studied our own solar system as well as disks of
successful observations obtained in ~23,500 h gas and dust around stars providing information
of science observing (exceeding the nominal mis- and comparisons with the global properties of
sion by ~20 %), covering almost 10 % of the sky, exosolar systems around nearby stars. Herschel
to the worldwide astronomical community. In has already shown that it will have large impact
addition, there are also over 3,000 science cali- on research in all these fields.
bration observations from about 1,700 h of
observing currently publicly available; this num- The Galactic Plane
ber will increase. Herschel has performed a complete 360 photo-
metric survey of the entire galactic plane in five
bands from 70 to 500 mm. Together with ancillary
Key Research Findings data at other wavelengths, this enormous dataset
will enable astronomers to address a number of
The “cool universe” fundamental questions regarding the global
Herschel is all about the “cool universe”; it is the workings of our Galaxy, in particular with respect
first and only space facility dedicated to the FIR/ to (high-mass) star formation and the resulting
sub-mm part of the spectrum. Roughly half the feedback on the ISM environment, see Fig. 2.
energy emitted in the universe since the forma- Dramatic structure on all scales is clearly visible.
tion of the cosmic microwave background First attempts to derive global star formation
(CMB) has been absorbed by dust in the ▶ inter- rates are underway, with the objective to con-
stellar medium (ISM) in our ▶ Milky Way and struct a new 3D model of the Galaxy, mapping
other galaxies and emitted at the longer Herschel the essential critical parameters like column den-
wavelengths appropriate to the ISM tempera- sity thresholds, rate, and efficiency of star forma-
tures. The integrated spectrum of the reradiated tion in the Galaxy.
Herschel Mission,
Fig. 3 An area in the
stellar nursery in the
constellation of Aquila
(ESA/SPIRE &
PACS/P. André)
A Stellar Nursery dimethyl ether, hydrogen cyanide, sulfur oxide,

Herschel’s view of a stellar nursery around 1,000 sulfur dioxide, and their ▶ isotope analogues.
▶ light-years away in the constellation Aquila This spectrum demonstrates the spectral richness
(the Eagle) is shown in Fig. 3. This cloud, of regions of star and planet formation; the com-
65 light-years across, is so shrouded in dust that plete survey has more than 20,000 lines and harbors
no infrared satellite has been able to see into it the promise of a deep understanding of the
until now. Thanks to Herschel’s greater sensitiv- astrochemistry of the region.
ity at the longest infrared wavelengths, astrono-
mers have their first picture inside this cloud.
Using Herschel’s PACS and SPIRE instruments Water
at the same time, the image shows two bright
regions where large newborn stars are causing A special talent of Herschel is its ability to
hydrogen gas to shine. Embedded in the dusty observe water in gas form. By a combination of
filaments are 700 condensations of dust and gas PACS and HIFI spectroscopy of Jupiter, astron-
that will eventually become stars. Astronomers omers have found direct proof that the bulk of
estimate that about 100 are “▶ protostars,” celes- the water currently present in Jupiter’s strato-
tial objects in the final stages of formation. Each sphere was delivered by comet Shoemaker-
one just needs to ignite nuclear fusion in its core Levy 9, which struck the planet in 1994. The
to become a true star. The other 600 objects are PACS observations in a grid of 25 points across
not developed sufficiently to be called protostars, the disk of Jupiter showed that the water is pre-
but eventually they will become another genera- dominantly confined to the southern hemi-
tion of stars. sphere, which is where the comet struck.
The HIFI observations indicate that the bulk
Astrochemistry Revealed of water is confined to pressures lower than
A part of a Herschel/HIFI spectral scan is shown in 2 millibars, corresponding to high stratospheric
Fig. 4. The observation is toward the Orion Nebula, altitudes – well above the uppermost part of the
a relatively nearby star-forming region, the troposphere which effectively acts as a “cold
“sword” in the constellation of Orion. A character- trap.” For these reasons, the water in the strato-
istic feature is the spectral richness: among the sphere is inferred to have been delivered from an
organic molecules identified in this spectrum are external source, identified as the 1994 comet
water, carbon monoxide, formaldehyde, methanol, impact (Fig. 5).
Herschel Mission, Fig. 4 A part of a Herschel/HIFI spectral scan (the white curve) overlaid on a background Spitzer
Space Telescope image (ESA/HIFI/HEXOS/E. Bergin)
Future Directions
“The Universe Explored by Herschel” sympo-

sium was held in October 2013 with ~360 partic-
ipants making in excess of 100 oral and ~200
poster presentations, providing the present status
and demonstrating that astronomers will be work-
ing with Herschel data for many years to come.
While the end of in-flight operations has been
a major mission milestone, at the same time in the
beginning of the postoperation phase, the Her-
schel scientific return in the form of rate of
refereed scientific publications has never been
higher and is still on the increase!
Herschel Mission, Fig. 5 The distribution of water in See Also

the stratosphere of Jupiter. White and cyan indicate highest
concentration of water, and blue indicates lesser amounts.
The map has been superimposed over an image of Jupiter
▶ Infrared Space Observatory
taken at visible wavelengths with the Hubble Space Tele- ▶ Interstellar Medium
scope (ESA/Herschel/T. Cavalié et al.; Jupiter image: ▶ Lagrangian Points
NASA/ESA/Reta Beebe (New Mexico State University)) ▶ Molecular Cloud
Hertzsprung-Russell Diagram 1099
▶ Molecules in Space William Herschel had only one child, John

▶ Protostars Frederick William Herschel (1792–1871), also
▶ Spitzer Space Telescope an astronomer who made important contributions
▶ Star Formation, Theory to stellar astronomy (namely star clusters and
▶ Stellar Evolution nebulae).
The European Space Agency’s Herschel
Space Observatory that was launched on May
14, 2009, was conceived to study the darkest
Herschel Publications List at: http://herschel.esac.esa.int/ and coldest parts of the Universe by the light of
ScientificPublications.shtml the far-infrared and submillimeter portions of the
Herschel Science Archive at: http://herschel.esac.esa.int/ spectrum.
Science_Archive.shtml
Herschel website at: http://herschel.esac.esa.int/
The Universe Explored by Herschel at: http://herschel.
esac.esa.int/TheUniverseExploredByHerschel.shtml References and Further Reading
Holmes R (2008) The age of wonder: how the romantic H

generation discovered the beauty and terror of science.
Pantheon, New York
Hoskin M (2013) William and Caroline Herschel: pio-
Herschel, William neers in late 18-th century astronomy. Springer, Berlin
Portugal
Hertzsprung-Russell Diagram
History Thierry Montmerle1 and Sylvia Ekström2

1
Institut d’Astrophysique de Paris, CNRS/
William Herschel (1738–1822) was a German- Université Paris 6, Paris, France
2
born musician and, most famously, astronomer, Observatoire Astronomique de l’Université de
who spent most of his life in Britain. He discov- Genève, Faculté des Sciences, Université de
ered Uranus in 1781, thereby doubling the size Genève, Versoix, Switzerland
of the known solar system. Other important
astronomical discoveries concerned the atmo-
sphere of Venus and others planets of the Solar Keywords
system: between 1787 and 1789 he discovered
Mimas and Enceladus, Saturn’s sixth and sev- Effective temperature; Luminosity; Stars; Stellar
enth satellites, and in 1787, he identified the first evolution
two satellites of Uranus, Oberon and Titania.
With the assistance of his sister Caroline
Herschel (1750–1848), an important astronomer Synonyms
by herself, whose most significant contribution
to astronomy was the discovery of several Color-magnitude diagram; HR diagram;
comets, he worked on a catalog of nebulae and Luminosity-temperature diagram
star clusters (the observational reductions were
primarily made by Caroline). As an astrobiolo-
gist, it is interesting to note that Herschel Acronyms
believed it likely that the Moon and planets
harbored life. HRD
1100 Hertzsprung-Russell Diagram
Definition 20–30 % in the case of young stars), and (2) to

convert from spectral types to temperatures,
The Hertzsprung-Russell diagram plots the which can be done with models of stellar photo-
▶ magnitude and color of stars, which are spheres. For “simple” stars like the Sun, the tem-
converted via theoretical models and distances perature determination is very precise (<1 %),
into stellar luminosities and surface (“effective”) but for more complex spectra, like the T Tauri
temperatures. Comparison with theoretical tracks, stars which have a circumstellar disk, the uncer-
the “paths” in the diagram corresponding to tainty may reach 10–20 % or more.
changes in observed properties as the star evolves,
then allows the determination of the mass or age of
a star or cluster. Note in the following the astro- Key Research Findings
physical use of the term “burning” to signify
nuclear fusion (e.g., of hydrogen to helium). Populated Regions
Since the beginning of the twentieth century, When the HRD is used with a sample of observed
astronomers have been using the “Hertzsprung- stars, it gives statistical information about the
Russell diagram” (from the name of its discov- surface characteristics of stars, defining specific
erers, in the early twentieth century; hereafter regions. Each region can be associated to a par-
HRD) to classify stars and understand their evo- ticular evolutionary phase by the knowledge of
lution, from their formation to their very late stellar evolution and theoretical tracks (see
stages. below). There are many interesting regions stand-
ing out in the diagram. Among them, one can
mention the following ones.
Overview
Main Sequence
When plotted in the HRD, the stars do not appear The great majority of the stars lie in the diagonal,
randomly distributed (see Fig. 1). There are some the main sequence (MS), which represents the
regions that are more populated and others that zone where stars convert hydrogen into helium,
are almost empty. There is an evident relation i.e., the first and longest phase of the nuclear life
between the density of a region and the duration of stars. This phase can last billions of years
of the ▶ stellar evolution phase that occurs in that for a star like the Sun. Since stars follow a
region. During their evolution, stars pass from mass-luminosity relation, with L/M3 during the
long-lasting hydrostatic burning (nuclear fusion) MS, the location of a star on the MS allows an
phases to quick structural readjustments. While it estimation of its mass. After this phase, the fusion
is easy to observe the stars in regions where the of hydrogen goes on in a shell, while the core
hydrostatic burnings occur, very few stars can be contracts and the envelope expands: the stars
observed during their readjustment phases. cross rapidly the HRD and become red giants or
supergiants.
Basic Methodology Giants and Supergiants

The giants and supergiants regions mark the loca-
Observationally, astronomers first determine the tion of the HRD where hydrostatic helium burn-
magnitude and ▶ spectral type of the stars. These ing occurs.
numbers are then transformed into luminosity Stars between 0.8 and 2 M (solar masses)
and temperature, which are physical quantities leave the MS to climb on the red giant branch
that can be compared with models. This assumes until they undergo a flash which ignites helium
the capability (1) to know the distance (to convert fusion in their core. After the flash, they settle on
magnitudes into luminosities), which is deter- the horizontal branch (giants region) where they
mined by various methods (to an accuracy of burn helium quietly. Between 2 and 10 M, the
Hertzsprung-Russell Diagram, Fig. 1 Hertzsprung- significant regions are indicated in gray. The duration of
Russell diagram (Courtesy Javier Garcı́a Nombela, art- the main sequence for the different masses is indicated in
eres.net). The observational units are in yellow (absolute red. Lines of isoradius are sketched in green. The names of
magnitude versus spectral class) and the theoretical ones some known stars are indicated
in white (luminosity versus effective temperature). Some
core helium burning starts smoothly, following This region is occupied by stars of different
the same path in the HRD. Above 10 M, most of types, among which two are commonly used as
the central helium burning occurs in the red distance calibrators:
supergiants region.
After central helium exhaustion, stars between 1. RR Lyrae, which are old giants of low
2 and 10 M populate the asymptotic giant ▶ metallicity on the horizontal branch
branch, where an unstable double shell burning 2. Classical Cepheids, which are helium burning
(a shell of helium burning and another of hydro- stars between 4 and 12 M
gen burning) drives thermal pulsations.
The distance calibration is based on the atten-
Instability Strip uation from the absolute magnitude (which rep-
The instability strip marks a region in the HRD resents the intrinsic stellar luminosity) to the
where stars become unstable to radial pulsations. apparent magnitude (reflecting the observed
Hertzsprung-Russell
Diagram, −12
log L/L⊙ M bol
Fig. 2 Theoretical tracks 120 M⊙
of stellar evolution in the 85 M⊙
Hertzsprung-Russell
6 60 M⊙
diagram (From Schaller −10
et al. 1992), showing 40 M⊙
luminosity (relative to the
Sun) (scale on the left) and
bolometric magnitude −8
(Mbol, scale of the right) 5 25 M⊙
versus effective 20 M⊙
temperature. The shaded
area represents the regions 15 M⊙ −6
of H and He burning. 12 M⊙
4
Stellar mass indicated
along the main 9 M⊙
sequence line −4
7 M⊙
3
5 M⊙
−2
4 M⊙
2 3 M⊙ 0
2.5 M⊙
Y = 0.300 2 M⊙
1 Z = 0.020 +2
1.7 M⊙
1.5 M⊙
1.25 M⊙ +4
0 1 M⊙
0.9 M⊙
04 07 09 B0 B1 B2 B3 B5 B8 A0 A3 A7 F1 F8 G2 0.8 M⊙ +6
4.8 4.7 4.6 4.5 4.4 4.3 4.2 4.1 4.0 3.9 3.8 3.7 3.6
log Teff
brightness; Hertzsprung 1913). While in the White Dwarf Location

normal case, it is extremely difficult to deter- At the end of the evolution, stars below 10 M
mine the absolute magnitude of a star, RR expel their envelope as a planetary nebula and
Lyrae and Cepheids present, respectively, a settle on the white dwarf branch, where they
metallicity-luminosity and a period-luminosity progressively cool.
relation that have long been thought to offer an
easy knowledge of their absolute magnitude,
since the periods and metallicities can be mea- Theoretical Tracks
sured. It is now known that those relations are When theoretical tracks are plotted in the HRD,
more complex and less direct than previously they offer an evolutionary link between the dif-
considered (see Sandage and Tammann 2006, ferent populations by giving the time sequence of
and references therein), and their use as distance the surface characteristics by which the stars pass
calibrators must be considered with caution. throughout their evolution.
Hertzsprung-Russell Diagram, Fig. 3 Hertzsprung- temperatures (Teff) are also indicated. Most stars appear
Russell diagram of 934 low-mass members of the young to be younger than a few million years, with masses from
Orion Nebula Cluster (Hillenbrand 1997) and theoretical 0.1 M or less, up to a few M‘ (higher masses are located
pre-main sequence tracks, labeled in masses and ages outside of this diagram) (From Montmerle et al. 2006)
(dotted lines). The spectral types corresponding to surface
An example of a theoretical HRD is given in successive nuclear reaction networks driving

Fig. 2 (Schaller et al. 1992). It plots the evolution important changes in the overall stellar structure.
of the stars from the central hydrogen ignition to Before the main sequence, however, the situ-
the end of helium burning. For low- and ation is entirely different, since the stars slowly
intermediate-mass stars, that is the end of the shrink as the radiation generated by gravitational
evolution, while for massive stars (above contraction is evacuated in the form of light. In
10 M), the nucleosynthetic activity continues other words, the energy of “pre-main sequence”
in the core, up to silicon burning. However, stars (protostars and T Tauri stars) is not nuclear
after helium exhaustion, the evolution of massive in origin but is drawn only from gravitation. This
stars accelerates significantly so the core and the “simplification” explains that, as early as 1966
envelope decouple: their external properties (when only low-power computers were avail-
remain almost identical throughout the advanced able!), theoretical models of pre-main sequence
phases up to the time of the supernova explosion. evolution could be devised by Hayashi and his
collaborators in Japan.
Before the Main Sequence: Hayashi Tracks This early work on “Hayashi” evolutionary
From the main sequence onwards, the energy tracks in the HRD distinguished two main phases
output is always thermonuclear in origin, with which are still used as a reference today:
Hertzsprung-Russell −8 AGE
Diagram, Fig. 4 Color- YR
magnitude diagram of MV NGC 2362 1.0 X 106
several galactic clusters h + X Perse
(From Sandage 1958). The −6 2.0 X 106
h + X Perse
age of the turn-off from the
main sequence is indicated
on the right axis. −4
Historically, this is the first 6.5 X 106
compilation of this type. M3
The shortest ages (less than Pletades
a few 10 million years) are −2 M41
2.8 X 107
wrong, because the “pre- M41 M11 M11
main sequence”
evolutionary phases were
−0
not known at the time Como H+P 1.6 X 108
Hyades M67
Proesepe M3
NGC 752 752

2
1.2 X 109
M67
4
Sun 7.1 X 109
6
2.9 X 1010
8
−4 0 .4 .8 1.2 1.6 2.0
B-V
1. The “convective” phase, in which the stars are early T Tauri stage (when the young Sun starts
fully convective and evolve essentially iso- to be optically visible). Then the radiative phase
thermally (i.e., at the same temperature, so lasts from 10 million to 100 million years at a
that the theoretical HRD tracks as a function roughly constant luminosity (1 L), until the
of mass are all vertical) thermonuclear reactions transforming hydrogen
2. The “radiative” phase, in which a radiative core into helium start. This marks the beginning of
develops inside the star, which then becomes the main sequence, already described, on which
hotter but evolves at an almost constant lumi- the Sun has been for 4.5 billion years and still
nosity (following the contraction in radius) is today.
Figure 3 illustrates pre-main sequence evolu-

tionary tracks, running through observational Applications
points for the young Orion nebula cluster: each
point represents a star, ordered by luminosity Once located in the HRD, observed stars can be
and temperature. For a star like the Sun, the compared to theoretical tracks. From such a con-
convective phase lasts about 10 million years, frontation, one can evaluate the mass and evolu-
during which the temperature is held almost tionary status of the star.
constant, around 4,600 K, and the luminosity The HRD can also be used to plot a whole
drops by a factor of 20 with respect to the cluster (see Fig. 3 for a young cluster and Fig. 4
Hesperian 1105
for an evolved cluster). The graph obtained will ▶ Pre-main-Sequence Star

be then compared to theoretical ▶ isochrones ▶ Spectral Type
(location of stars with different masses at a ▶ Star Formation, Theory
given age), so the age of the cluster can be ▶ Stellar Evolution
determined. ▶ Supernova
The comparison between observed stars and
theoretical tracks is neither straightforward nor
simple. As mentioned above, the comparison References and Further Reading
between observation and theory implies a con-
version from observational units (color and mag- Bertout C (1989) T tauri stars: wild as dust. Annu Rev
nitude) to theoretical ones (effective temperature Astron Astrophys 27:351–395
Hayashi C (1966) Evolution of protostars. Annu Rev
and luminosity), with some uncertainties in the Astron Astrophys 4:171–192
conversion. Some observed stars may be Hertzsprung E (1913) Über die räumliche Verteilung der
unresolved binaries, giving access only to the Veränderlichen vom d Cephei-Typus. Astron Nachr
addition of their magnitude and to the mean of 196:201–210
their color index. Some stellar parameters (as the
Hillenbrand L (1997) Astron J 113:1733–1768 H
Montmerle T et al (2006) From suns to life: a chronolog-
chemical composition or the rotation rate, e.g.) ical approach to the history of life on earth 3. Solar
can modify the location in the HRD of the theo- system formation and early evolution: the first 100 mil-
retical tracks. A good knowledge of the observed lion years. Earth Moon Planets 98:39–95
Sandage A (1958) The color-magnitude diagrams of
object and an adequate choice of the stellar galactic and globular clusters and their interpretation
models are needed to get valuable information as age groups. Ric Astron 5:41–68
from such a comparison. Note that some physical Sandage A, Tammann GA (2006) Absolute magnitude
inputs in the models are still uncertain (treatment calibrations of population I and II cepheids and other
pulsating variables in the instability strip of the
of convection, mass loss prescription, magnetic Hertzsprung-Russell diagram. Annu Rev Astron
fields) and can alter the tracks. Astrophys 44:93–140
Schaller G, Schaerer D, Meynet G, Maeder A (1992) New
grids of stellar models from 0.8 to 120 solar masses at
Z = 0.020 and Z = 0.001. J Astron Astrophys
Future Directions 96:269–331, Supplement Series
Stahler S, Palla F (2005) The formation of stars. Wiley
The stellar evolutionary tracks are established for Interscience, New York. ISBN 3-527-40559-3
a fixed mass sufficiently precisely that one can
have an estimate of the mass and age of a star. But
a major phenomenon still defies modelers. This is
mass loss and its interplay with evolution, for Hesperian
which only crude theories exist. In the case of
T Tauri stars, especially in the early stages, accre- Ernst Hauber
tion (from a circumstellar disk) is important but is Deutsches Zentrum f€ur Luft- und Raumfahrt
taken only phenomenologically into account in (DLR) e.V., Institut f€ur Planetenforschung,
the evolutionary models. Berlin, Germany
See Also Definition
▶ Isochrone The middle of three systems (of time-

▶ Magnitude stratigraphic units) or periods (the chronological
▶ Main Sequence, Star equivalents to systems) in the Martian strati-
▶ Metallicity graphic scheme, named after the region of
1106 Heterocycle
Hesperia Planum (Hesperia: “The Occident”; see

U.S. Geological Survey Gazetteer of Planetary Heterotroph
Nomenclature). Depending on the different
models to determine absolute ages on planetary Felipe Gomez
surfaces by crater statistics, the ▶ Hesperian Centro de Astrobiologı́a (CSIC/INTA), Instituto
began 3.8 billion years ago and ended at some Nacional de Técnica Aeroespacial, Torrejón de
point between 3.55 and 1.8 billion years ago. Ardoz, Madrid, Spain
See Also
Definition
▶ Amazonian
A heterotroph is any organism that needs
organic compounds as a ▶ carbon source for
▶ Crater, Impact
the synthesis of its own cellular components.
▶ Mars
According to their source of energy, organisms
▶ Mars Stratigraphy
can be classified as photoheterotrophs or
▶ Noachian
chemoorganotrophs. Chemoorganotrophs often
use reduced organic compounds as a source of
both energy and carbon. Mixotrophs using inor-
ganic compounds as an energy source and
Heterocycle reduced organic compounds as a carbon source
are an exception.
Shin Miyakawa
See Also
Definition
A heterocycle is an organic compound which
▶ Biosynthesis
contains within a ring structure, which can be
▶ Carbon Source
aliphatic or aromatic, at least one element other
▶ Chemoorganotroph
than carbon, such as nitrogen, oxygen, or sulfur.
▶ Phototroph
Examples are purines, pyrimidines, imidazoles,
and quinolines. These structures may exist in
PAHs in meteorites and dusts and in ▶ Titan
hazes. In prebiotic experiments, these com-
pounds can be formed by electric discharges and Heterotrophic Hypothesis
proton irradiation of gas mixtures containing car-
bon, nitrogen, oxygen, and hydrogen, such as Robert Pascal
CH4-N2-H2O. Institut des Biomolécules Max Mousseron
CC1706, Université de Montpellier II,
Montpellier, France
See Also
▶ Nucleic Acid Base Synonyms

▶ PAH
▶ Titan Prebiotic soup hypothesis
Hexamethylenetetramine 1107
Definition Miller SL (1953) Production of amino acids under possi-

ble primitive earth conditions. Science 117(3046):528
Oparin AI (1924) The origin of life. Moscow Worker,
According to the heterotrophic hypothesis for the Moscow
origin of life, early organisms depended on abiot- Urey HC (1952) On the early chemical history of the earth
ically synthesized organic molecules for their and the origin of life. Proc Natl Acad Sci U S A 38:
structural components and as an energy source. 351–363
The hypothesis is usually considered in connection
with abiotic organic matter of atmospheric origin,
but is also consistent with extraterrestrial inputs
(▶ meteorites, micrometeorites, comets) or hydro- Hexamethyleneamine
thermal synthesis. The dilution of organic matter
in the ocean is considered sufficient reason to rule ▶ Hexamethylenetetramine
out its relevance by many opponents.
History Hexamethylenetetramine H
In 1871, the mention of a “warm little pond” by Greg Springsteen

Darwin may be considered as the first mention of Furman University, Greenville, SC, USA
this hypothesis. Subsequently, it was more
clearly formulated by Oparin (1924), Haldane
(1929), and Urey (1952), ideas which ultimately Keywords
led to the Miller experiment (1953).
Ammonia; Condensation; Formaldehyde
See Also
▶ Abiotic Photosynthesis Synonyms

▶ Atmosphere Escape
▶ Atmosphere, Organic Synthesis 1,3,5,7-Tetraazatricyclo[3.3.1.13,7]decane;
▶ Chemical Evolution 11,3,5,7-Tetraazaadamantane; Formamine;
▶ Darwin’s Conception of the Origins of Life Hexamethyleneamine; Hexamine; HMT; Methe-
▶ Earth’s Atmosphere, Origin and Evolution of namine; Urotropine
▶ Extraterrestrial Delivery of Organic Compounds
▶ Metabolism
▶ Oparin’s Conception of Origins of Life Definition
▶ Organic Material Inventory
▶ Origin of Life An adamantane-like heterocycle (C6H12N4)
▶ Photochemistry, Atmospheric formed from the condensation of six ▶ formalde-
▶ Prebiotic Chemistry hyde and four ▶ ammonia molecules.
▶ Primordial Soup
▶ Urey’s Conception of Origins of Life
Overview
References and Further Reading Hexamethylenetetramine (HMT) is a condensa-
tion product formed readily from gaseous, aque-
Haldane JBS (1929/1967) “The origin of life”. The ratio-
nalist annual. Reprinted as an appendix In: Bernal JD ous, or organic solutions of ammonia and
(ed) The origin of life. Weidenfeld & Nicolson, London formaldehyde. It was first synthesized and
1108 Hexamine
characterized by the Russian chemist Aleksandr References and Further Reading

Butlerov (who also discovered the ▶ formose
reaction) in 1859 and was the first organic com- Bernstein MP, Sandford SA, Allamondola LJ, Chang
S (1994) J Phys Chem 98(47):12206–12210
pound with a solved X-ray crystal structure. HMT
Blazevic N, Koldbah D, Belin B, Sunjic V, Kajfez
has a molecular formula of C6H12N4 and is gen- F (1979) Hexamethylenetetramine: a versatile reagent
erated in a highly exothermic condensation in organic synthesis. Synthesis 3:161–176
(55 kcal/mol) from six formaldehyde and four Cleaves HJ (2008) The prebiotic geochemistry of formal-
dehyde. Precambrian Res 164:111–118
ammonia molecules with the loss of six waters.
Dondi D, Meril D, Pretali L, Fagnoni M, Albini A,
The exothermicity results from the transforma- Serpone N (2007) Prebiotic chemistry: chemical evo-
tion of six pi bonds into six sigma bonds. It is a lution of organics on the primitive Earth under simu-
white crystalline powder with a sublimation tem- lated prebiotic conditions. Photochem Photobiol Sci
11:1210–1217
perature of 285–295 C and an adamantane-like
chemical structure with tetrahedral symmetry. It
is soluble in chloroform, ethanol, and water
(850 g/L at 25 C) and insoluble in diethyl ether. Hexamine
It is commercially produced by passing gas-
eous ammonia into a concentrated solution of ▶ Hexamethylenetetramine
aqueous formaldehyde with subsequent removal
of water under reduced pressure. Solid HMT can
be more easily obtained on a small scale by pass-
ing gaseous ammonia into a solution of parafor- HGT
maldehyde dissolved in refluxing toluene with
concomitant water removal (i.e., a Dean- ▶ Lateral Gene Transfer
Stark trap).
HMT is stable in neutral and basic aqueous
media, with an equilibrium constant approaching High Accuracy Radial-Velocity Planet
1010. In acidic media, however, it is hydrolyzed Searcher
to methanimine (H2CNH), formaldehyde, and
ammonia. Synthetically, HMT is useful in the ▶ HARPS
alkylation of primary amines (Delepine reaction)
and in the generation of amino alcohols by reac-
tion with oxiranes.
HMT is produced as the primary product High Resolution Echelle
(35 % yield after 45 days at rt) from the UV Spectrometer
irradiation of methanol-ammonia-water mixtures
and has been synthesized in laboratory experi- ▶ HIRES
ments that approximate conditions in the inter-
stellar medium. The spectral properties of HMT
and its photolysis products (namely, the High-Efficiency Particulate
2,160 cm1 absorption) indicate that it may com- Absorbing Filters
prise a portion of interstellar ice grains and icy
satellites. ▶ HEPA Filters
See Also
High-Efficiency Particulate Air Filters
▶ Ammonia
▶ Formaldehyde ▶ HEPA Filters
HII Region 1109
Gravitational collapse of the Fe core turns into a

High-Efficiency Particulate Arresting supernova explosion, ejecting most of the mass in
Filters the form of heavy elements. They are the major
nucleosynthesis sites in the universe, and they leave
▶ HEPA Filters behind either a ▶ neutron star or a ▶ black hole.
See Also
High-Energy Photons
▶ Black Hole
▶ Gamma Rays
▶ Neutron Star
▶ Star
High-Energy Radiation ▶ Nucleosynthesis, Stellar
▶ Supernova
▶ Gamma Rays ▶ Zero Age Main Sequence H
Highland Region High-Performance Liquid

Chromatography
▶ Terra, Terrae
▶ HPLC
High-Magnesium Granodiorite
High-Pressure Liquid
▶ Sanukitoid Chromatography
▶ HPLC
High-Mass Star
Nikos Prantzos HII Region

Daniel Rouan
Definition Meudon, France
Stars with an initial mass on the ▶ zero age main

sequence above 8 M are considered to be high- Synonyms
mass stars. They evolve in a short timescale
(<25 Myr) by nuclear fusion of successively all H+ region
available nuclear fuels, up to 56Fe. The most mas-
sive of them suffer heavy mass losses through
stellar winds, revealing their nuclear-processed Definition
layers (He rich or C rich). At the end of their life,
they have developed an “onion skin” structure, with An H II region is an astrophysical object belong-
heavier elements lying toward the center. ing to the ▶ interstellar medium and consisting in
1110 Hill Radius/Sphere
HII Region, Fig. 1 The

Orion nebula is the
prototype H II region, that
is, a cloud of hydrogen in
the Galaxy, ionized by hot
stars in the vicinity. The red
color results from the
recombination of protons
and electrons producing a
cascade of energy with
emission of light
a cloud of hot ▶ plasma of protons and electrons in the radio domain. Compact H II regions des-
at about 10,000 K. The name stems from the old ignate the youngest ones. The Orion nebula, eas-
appellation H II for the ionized state of hydrogen ily seen with binoculars, is one archetype of an
(now H+), as opposed to the neutral state H I. The H II region (Fig. 1).
plasma is produced by photoionization of
the hydrogen in an interstellar cloud caused by
the ultraviolet radiation from nearby hot massive See Also
stars of ▶ spectral type O and B. The stars are
generally born in a giant ▶ molecular cloud and ▶ Interstellar Medium
ionize the gas around them that was not con-
densed into stars. The occasional recombination
of a proton and an electron produces a cascade in
the energy levels with emission of photons that Hill Radius/Sphere
explains why H II regions appear as luminous
nebulae. Their spectrum contains mainly strong Sean N. Raymond
H I recombination lines, especially the Ha line Laboratoire d’Astrophysique de Bordeaux,
responsible for the reddish color, as well as for- CNRS, Universite de Bordeaux, France
bidden lines (transitions from metastable states,
denoted in astronomy by square brackets) such as
those from ionized oxygen and nitrogen ([O III], Synonyms
[O II], [N II]). The plasma is also a source of free-
free (bremsstrahlung) emission mainly detected Roche radius
Hipparcos 1111
Definition See Also
The region around a planet within which ▶ Exoplanets, Discovery

the planet’s gravity dominates that of the star it ▶ Orbit
orbits is called the Hill sphere, whose radius is
called the Hill radius. The Hill radius is given by
1=3
RHill ¼ a mp =3Mstar , where a is the orbital
distance of the planet from the star, mp is the Hipparcos
planet’s mass, and Mstar is the stellar mass.
The Earth’s Hill radius is roughly 0.01 AU, Michel Viso
and Jupiter’s is about 0.35 AU. The Hill sphere CNES/DSP/SME, Vétérinaire/DVM, Astro/
is the same as the Roche sphere or Roche Exobiology, Paris Cedex 1, France
lobe, but is not to be confused with the Roche
limit.
Definition
H
See Also In 1981, ESA approved a mission covering, for
the Agency, a new astronomical field. Hipparcos
▶ Planet Formation (originally an acronym for High Precision Paral-
lax Collecting Satellite) was developed to deter-
mine the positions, distances, and proper motions
of more than 100 000 stars with an accuracy
improved by a factor of 10–100 compared to
Hill/Lagrange Stability ground-based observations. In the process, it col-
lected also data on their variability and identified
Rory Barnes and characterized binary systems.
Astronomy Department, University of Hipparcos, a 500 kg spacecraft, was launched
Washington, Seattle, WA, USA into geostationary transfer orbit by an Ariane-4
launcher on 8 August 1989. However, the apogee
boost motor did not fire, and it could not reach its
planned geostationary orbit. Operations were
Definition conducted from a highly elliptical orbit with a
period of around 10 h.
In planetary dynamics, stability refers to an
aspect of the orbital motion that remains con-
stant in time, for example, that all planets Overview
remain bound to their star. Hill stability
only requires that the ordering of the orbiting By November 1989, Hipparcos started its opera-
bodies remain constant, meaning that the outer- tional mission. The observations lasted 3.5 years,
most planet may leave the system. The advan- ending in August 1993. Processing and verifying
tage of considering Hill stability is that it can the data took another 3 years. The Hipparcos
be analytically treated. In contrast, Lagrange Catalogue, with 118 218 entries, was finalized
stability requires any and all orbital oscillations on 8 August 1996, surpassing all of the original
to remain bounded for all times. Lagrange sta- scientific goals. A larger but less accurate “Tycho
bility is, therefore, a type of Hill stability, but Catalogue,” derived from the star mappers of the
can only be assessed via N-body simulations. satellite’s attitude control system and incorporat-
Many planetary systems appear to lie near these ing 1 058 332 star positions, was also compiled.
boundaries. In 1997, the data set was published in 17 volumes.
1112 HIRES
The enlarged Tycho 2 Catalogue of 2.5 million radial-velocity orbits for stars hosting exoplanets
stars was published in 2000. in order to derive planetary masses. Because the
Hipparcos was the first space mission dedicated input is a slit, an iodine gas absorption cell is
to measuring the positions of the stars. Beyond its introduced into the stellar beam in order to cali-
impact on basic stellar physics and on galactic brate the instrumental profile and wavelength scale
structure and dynamics, Hipparcos helped to pre- of each stellar observation. The data analysis
dict the impacts of ▶ Comet Shoemaker-Levy 9 on requires also a high-quality template observation
▶ Jupiter, identified stars that will pass close to the of each star without the iodine and a fundamental
Sun, established the distances of stars possessing calibration of the spectrum of the absorption cell at
planets even though these planets were discovered very high spectral resolution. HIRES has achieved
only after the end of the mission, identified a group a performance approaching velocities as low as
of stars that were captured by our Galaxy when it 1 m/s. HIRES has contributed to the discovery of
was young, fundamentally improved the cosmic several hundred extrasolar planets including the
distance scale (making the universe bigger and first super-Earth, GJ 876 d, several potentially
younger), and confirmed Einstein’s prediction of habitable planets around stars such as GJ 667C
the effect of gravity on starlight. and Gliese 581, and the confirmation of Kepler
A new ESA mission for astrometry, Gaia, was 78b, an Earth-sized planet detected by the Kepler
launched using a Soyouz Fregat in December space telescope. HIRES and ▶ HARPS on the
2013 from Kuru in French Guiana. 3.6-m telescope at the European Southern Obser-
vatory on La Silla are now the premier instruments
for determining the masses of small planets.
See Also
▶ Gaia Mission See Also
▶ HARPS
HIRES ▶ Radial-Velocity Planets

1
Cambridge, MA, USA Histidine
2
Manoa, Honolulu, Hawaii, HI, USA Henderson James (Jim) Cleaves II
Synonyms Japan
High Resolution Echelle Spectrometer Blue Marble Space Institute of Science,
Washington, DC, USA
Definition of Technology, Atlanta, GA, USA
HIRES (High Resolution Echelle Spectrometer) is

a high-resolution spectrograph on the Keck Definition
I telescope in Hawaii. It was originally designed
as a general-purpose instrument, but one of its Histidine is one of the 20 ribosomally encoded
main applications has been the determination of protein ▶ amino acids. It was first isolated by the
Homeostasis 1113
German physician and biochemist Albrecht Kossel

in 1896. It is distinguished by having a side chain HO2
with an imidazole ring, which has a pK a of approx-
imately 6. Histidine has an isoelectric point (pI) of ▶ Water, Related Interstellar Radicals and Ions
7.6; thus, at typical intracellular pH values, rela-
tively small shifts in pH can greatly change its
average charge. When protonated, the imidazole HOCH2CH2OH
ring is positively charged, with the positive charge
distributed between both ring nitrogen atoms, ▶ Ethylene Glycol
which can be represented by two resonance struc-
tures. The imidazole ring of histidine is aromatic at
all pH values as it contains six p electrons, is a
common coordinating ligand in metalloproteins,
HOCH2CHO
and is a part of the catalytic sites of many enzymes.
▶ Glycolaldehyde
Histidine has not been identified in carbona-
ceous chondrites and is difficult to be formed in H
Miller-type prebiotic synthesis experiments,
while imidazole can be easily obtained in them. HOCH2COCH2OH
▶ Dihydroxyacetone
See Also
▶ Amino Acid Holism

▶ Enzyme
▶ Heterocycle ▶ Vitalism
HMT Holophyletic
▶ Monophyletic
▶ Hexamethylenetetramine
Homeostasis
HNC
Alvaro Moreno1 and Kepa Ruiz-Mirazo2
▶ Hydrogen Isocyanide 1
Departamento de Lógica y Filosofı́a de la
Ciencia, Universidad del Paı́s Vasco, San
Sebastián, Spain
2
Department of Logic and Philosophy of Science,
HNCHCN
FICE, UPV-EHU, Biophysics Research Unit
(CSIC – UPV/EHU), Donostia, San Sebastián,
▶ Cyanomethanimine
Spain
Definition
HNCNH
Homeostasis consists in the maintenance of the
▶ Carbodiimide conditions in which a system is viable, despite
1114 Homochirality
external perturbations or differences with its constituent molecules are the same enantiomer.
environment. Homochirality is an important feature of terres-
trial biochemistry. All life on Earth is homochiral
(with rare exceptions); only L-amino acids are
encoded in proteins, and only D-sugars form the
History
backbones of DNA and RNA.
The French biologist Claude Bernard coined the
term in 1865 (from Greek homoios, “similar,”
Overview
and histemi, “standing still”), as the capacity of
living systems to maintain a stable, functionally
Homochirality is a unique feature of life on
adequate “internal milieu” through some mecha-
Earth (Castelvechhi 2007). Most biomolecules
nism of regulation.
occur in nonsuperimposable mirror image forms
or chiral isomers (enantiomers). The name chi-
ral derives from the Greek word for hand, and
See Also
biomolecules displaying a mirror symmetry are
either “left-handed” (L) or “right-handed” (D).
▶ Cell
Life favors one handedness: all known living
▶ Metabolism
organisms almost exclusively use L-amino
acids and D-sugars in their biochemistry, with
few exceptions. This preference extends to the
macromolecular level where biopolymers are
Homochirality assembled from chiral subunits such that all
monomers possess the same chirality: one
Sara Imari Walker finds only L-amino acids in proteins and only
School of Earth and Space Exploration, Arizona D-sugars in ▶ DNA and ▶ RNA, with no trace
State University, Tempe, AZ, USA of the “unnatural” D-amino acids and L-sugars
in either type of polymer chain (Bonner 1996).
The preference for only one molecular handed-
Keywords ness is referred to as homochirality and is essen-
tial to the proper folding of biopolymers and the
Biochirality; Enantiomeric excess; functioning of most biochemical processes in
Biosignatures; Chiral selection; Chirobiogenesis; extant life. Despite the prominence of
Mirror symmetry homochirality in biochemistry, the origin of
biological homochirality is not known and pre-
sents one of the longest standing mysteries in
Synonyms studies of the origin of life (Bada 1995). Labo-
ratory syntheses of amino acids and sugars gen-
100 % enantiomeric excess; Enantiopure erally yield racemic mixtures (equal amounts of
both the L- and D- enantiomers) under
prebiotically relevant conditions (Mason
Definition 2000). Therefore, a chiral selection process
was required at some stage in the origin or
Homochirality refers to the property of a group of early evolution of life on Earth. Various theo-
molecules composed of chiral units (enantio- ries have been proposed to explain how
mers). A substance is homochiral if all homochirality might have first emerged. Most
Homology 1115
theories require a chiral amplification mecha- ▶ Meteorite, Murchison

nism which acts to enhance a small initial asym- ▶ Polarized Light and Homochirality
metry. Small excesses of L-amino acids have ▶ Racemic Mixture
been observed in meteoritic samples such as
from the Murchison meteorite (Cronin and
Pizarello 1997) and provide the only natural
example of molecular asymmetry measured Bada JL (1995) Origins of homochirality. Nature
outside of the biosphere. Other postulated 374:594–595
sources of an initial asymmetry which might Bailey J (2004) Astronomical sources of circularly polar-
have seeded life’s homochirality include the ized light and the origin of homochirality. Orig Life
electroweak interaction (Cline 2005); the influ- Blackmond D (2010) The origin of biological homochirality.
ence of an asymmetric environment, such as Cold Spring Harb Perspect Biol 2:a002147
certain mineral surfaces (Hazen 2004) or that Blackmond D, Klussmann M (2007) Spoilt for choice:
induced by circularly polarized light (Bailey assessing phase behavior models for the evolution of
homochirality. Chem Commun 39:3990–3996
2004); or random statistical fluctuation (Frank Bonner WA (1996) The quest for chirality. In: Cline D -
1953). All of these sources of asymmetry pro- (ed) Physical origin of homochirality in life. American
H
duce only a small enantiomeric excess, and Institute of Physics Press, New York, pp 17–49
some type of amplification mechanism is there- Castelvechhi D (2007) Alien pizza, anyone? Sci News
172:107–109
fore required to achieve the complete chiral Cline D (2005) On the physical origin of the homochirality
symmetry breaking as observed in living sys- of life. Eur Rev 13:49–59
tems (Plasson et al. 2007; Blackmond 2010). Cronin J, Pizarello S (1997) Enantiomeric excesses in
Amplification can occur through ▶ autocataly- meteoritic amino acids. Science 275:951–955
Frank F (1953) On spontaneous asymmetric synthesis.
sis, when one enantiomer acts as a catalyst for Biochim Biophys Acta 11:459–463
its own formation while simultaneously Hazen RM (2004) Chiral crystal faces of common rock-
suppressing the formation of the other enantio- forming minerals. In: Palyi G et al (eds) Progress in
mer (Frank 1953), or through crystallization biological chirality. Elsevier, New York, pp 137–151
Mason S (2000) Extraterrestrial handedness revisited.
processes (Viedma 2005) or phase behavior Orig Life Evol Biosph 30:435–437
(Blackmond and Klussmann 2007). However, Plasson R et al (2007) Emergence of homochirality in
no mechanism for the origin of life’s far-from equilibrium systems: mechanisms and role
homochirality has been conclusively deter- in prebiotic chemistry. Chirality 19:589–600
Viedma C (2005) Chiral symmetry breaking during crys-
mined. Due to the universal nature of life’s tallization: complete chiral purity induced by nonlinear
observed homochirality, the question of chiral- autocatalysis and recycling. Phys Rev Lett 94:065504
ity in potential extraterrestrial biochemistries is
pertinent to astrobiological searches for life.
Homochirality has therefore been suggested as
a potential biosignature. Homology
David Moreira
See Also Unité d’Ecologie, Systématique et Evolution
CNRS UMR8079, Université Paris-Sud 11, Paris,
▶ Achiral Orsay, Cedex, France
▶ Amino Acid
▶ Autocatalysis
▶ Chirality Keywords
▶ Enantiomeric Excess
▶ Enantiomers Common ancestry; Phylogeny; Similarity
1116 Homology
Definition that statements frequently seen in literature like

“high homology” or “percent homology” are
Homology, one of the key concepts in modern incorrect (“high similarity” and “percent simi-
evolutionary biology, refers to the similarity larity” would be the correct terms). Likewise,
between characteristics of organisms due to a homology does not imply the maintenance of
common origin from a ▶ common ancestor (i.e., identical functions in different organisms. In
shared ancestry). Homology can be applied to fact, homologous structures can evolve to carry
organs and anatomical structures but also to out different functions, such as the wings of bats
molecular characters such as DNA sequences. versus the arms of other mammals. Genes
When characters in different organisms are sim- derived from a common ancestral gene but car-
ilar but they do not share a common evolutionary rying out different functions are very common.
origin, they are called analogous (e.g., the wings For example, enzymes involved in cell metabo-
of birds and insects) and they are the product of lism in many species, such as the lactate dehy-
convergent evolution. The accurate distinction drogenase, are used in animals also as structural
between true homologous characters and analo- proteins called crystallins to build up the lenses
gous ones is an essential step for phylogenetic of eyes.
analyses. Although similarity is not a strict criterion to
determine homology because of potential prob-
lems of convergent evolution (e.g., integral
membrane proteins tend to accumulate hydro-
History
phobic amino acids, artificially increasing the
similarity of sequences), it is often used to sug-
The biological concept of homology was first
gest homology between DNA and protein
used by the French naturalist Étienne Geoffroy
sequences (Zuckerkandl and Pauling 1965).
Saint-Hilaire (1772–1844) to describe anatomi-
In those cases, probabilistic criteria are com-
cal similarities in animals (Saint-Hilaire 1818).
monly used to avoid considering as homologous
Organs of different animal species occupying the
sequences that resemble each other by chance.
same position and having similar connections
The correct identification of homologous
with other organs were defined as homologous.
sequences is an essential prerequisite for molec-
This approach allowed defining body plans that
ular phylogenetic analyses, to avoid the use of
were common to very diverse species despite
convergent characters that lead to the recon-
their apparent dissimilarity. Later on, homology
struction of artifactual trees. There are two
studies were carried out in other organisms
major types of homologous sequences:
and, during the twentieth century, also to molec-
orthologous (those that diverge following the
ular characters, including DNA and protein
divergence of species) and ▶ paralogous (those
sequences.
originated from gene duplication) ones (Fitch
2000).
Overview
Homology is often confounded with similarity, See Also

which is erroneous. Homology is an absolute
criterion: two characters (e.g., two genes) are ▶ Common Ancestor
homologous or not, namely, they derive from a ▶ Gene
common ancestral character (e.g., an ancestral ▶ Orthologous Gene
▶ gene) or not. This means that homology, in ▶ Paralogous Gene
contrast with similarity, cannot be quantified, so ▶ Phylogeny
Hoogsteen Pair 1117

Hoogsteen Pair
Fitch WM (2000) Homology a personal view on some of
the problems. Trends Genet 16:227–231
Graur D, Li W-H (1999) Fundamentals of molecular evo-
lution. Sinauer Associates, Sunderland Earth–Life Science Institute (ELSI), Tokyo
Mayr E (1978) Origin and history of some terms in sys- Institute of Technology, Meguro–ku, Tokyo,
tematic and evolutionary biology. Syst Zool 27:83–88 Japan
Mindell DP, Meyer A (2001) Homology evolving. Trends
Ecol Evol 16:434–440
Saint-Hilaire G (1818) Philosophie anatomique des USA
organes respiratoires sous le rapport de le détermina- Blue Marble Space Institute of Science,
tion et de l’identité de leurs pièces osseuses. Washington, DC, USA
Méquignon-Marvis, Paris
Zuckerkandl E, Pauling L (1965) Evolutionary divergence
and convergence in proteins. In: Bryson V, Vogel HJ of Technology, Atlanta, GA, USA
(eds) Evolving genes and proteins. Academic, New
York, pp 97–166
Definition H
Homolysis A Hoogsteen Pair is a non-canonical/non-

Watson-Crick hydrogen-bonded motif observed
Henderson James (Jim) Cleaves II in nucleic acids.
Earth–Life Science Institute (ELSI), Tokyo Two nucleosides from two different nucleic
Institute of Technology, Meguro–ku, Tokyo, Japan strands, or from distal regions of the same strand,
Institute for Advanced Study, Princeton, NJ, USA can be held together by Hoogsteen base pairing.
Blue Marble Space Institute of Science, Hoogsteen base-pairs often involve the N7 posi-
Washington, DC, USA tion of purine bases as H-bond acceptors and the
Center for Chemical Evolution, Georgia Institute C6 exocyclic amino group of adenine residues as
of Technology, Atlanta, GA, USA H-donors, which then bond with the Watson-
Crick surface of pyrimidine bases.
Karst Hoogsteen discovered the eponymous
Synonyms
motif after observing a crystal structure of a
nucleic acid complex in which analogues of ade-
Homolytic fission
nine and thymine formed base pairing motifs with
a different geometry from the Watson and Crick
Definition motif.
Hoogsteen pairs have considerably different
In chemistry, homolysis is the scission of a chemical geometries from Watson-Crick base pairs and
bond in a neutral molecule to give two free radicals. consequently distinct behavior. Hoogsteen base
pairs exist transiently in some DNA molecules in
A B ! A þB (1) thermal equilibrium with standard Watson–Crick
base pairs. They are also observed in protein-
Homolytic reactions can be induced by UV or DNA complexes and G-quadruplexes.
visible irradiation (photolysis) or by peroxides.
Homolytic Fission See Also
▶ Homolysis ▶ Hydrogen Bond

1118 HOOH
sulfur bacteria) and 3-methylhopanoids from Type

HOOH I aerobic methylotrophic bacteria.
Hopanes are observed in carbonate and shale
▶ Water, Related Interstellar Radicals and Ions sediments as old as 1.65 Ga. They are common
of, but not limited to, oceanic, lacustrine, hydro-
thermal environments throughout geological his-
tory but are markedly abundant in sediments from
some extreme depositional settings, such as the
Hopanes, Geological Record of Cretaceous oceanic anoxic events (OAEs). The
distribution of hopane structures in a kerogen, oil,
Jennifer Eigenbrode and bitumen often reflects the degree of thermal
NASA Goddard Space Flight Center, Greenbelt, maturity and correlates to relative carbonate
MD, USA verses clay content.
Definition See Also
Hopanes is a class of pentacyclic compounds ▶ Cyanobacteria

derived from bacteriohopanepolyols (hopanoids) ▶ Isoprenoids
via diagenetic degradation and thermal matura- ▶ Molecular Fossils
tion. The hopane structure is a saturated hydrocar-
bon (Fig. 1). Hopanes are molecular fossils of
bacterial lipids that serve to adjust membrane per-
meability and fluidity as well as other adaptation to Horizontal Branch
extreme habitats. Some hopanoids and derived
hopanes are regarded as specific to certain groups Nikos Prantzos
of bacteria, such as 2-methyl hopanoids from pho- Institut d’Astrophysique de Paris, Paris, France
tosynthetic bacteria (▶ cyanobacteria and purple
Definition
The horizontal branch (HB) is the region of the

Hertzsprung-Russell diagram occupied by
evolved stars – after the ▶ red-giant stage – of
low mass and low metallicity (abundance of ele-
ments heavier than He less than 0.1 that of the
Sun), during their central He-burning (He-fusion)
phase. The HB is observed in globular clusters,
whereas in open clusters (having approximately
solar ▶ metallicity), stars are confined to the bot-
Hopanes, Geological Record of, Fig. 1 Core structure tom of the red-giant branch during core He burn-
of hopanes composed of C-C bonds (lines) and saturated ing. Metallicity is the key factor explaining that
with hydrogen atoms (not shown). The key stereochemical difference (low-metallicity stars have lower
variants occur at C-17, C-21, and C-22 positions. Addi-
opacities and are hotter), but other factors
tional methylation is common at C-2 and C-3. Other
structural variants are missing methyl groups. R = H, (composition, age, rotation, envelope mass)
methyl, ethyl, n-propyl, n-butyl, or n-pentyl group appear to affect the morphology of the HB.
Hot Core 1119
See Also Overview
▶ Globular Cluster Hot molecular cores and hot corinos are charac-
▶ Hertzsprung-Russell Diagram terized physically by high gas densities
▶ Metallicity (>106 cm3) and elevated temperatures of both
▶ Open Cluster gas and dust (100–500 K). The gaseous chem-
▶ Red Giant ical composition is distinct from that of cold
molecular clouds, due to evaporation of ice man-
tles from dust grains, and contains many complex
organic molecules. The high temperatures are
produced by the luminosity of the protostar and
Horizontal Gene Transfer the protostellar accretion disk. Shock waves may
also play a chemical role in both heating the gas
▶ Lateral Gene Transfer and in ▶ sputtering icy grain mantles back into
the gas.
Hot cores are chemically rich and display the H
products of interstellar grain-surface chemistry in
the gas phase. For this reason, they can be used to
Hot Core probe catalytic processes on dust grains. Among
the species found in them are methanol, ethanol,
Steven B. Charnley dimethyl ether, ethyl methyl ethyl, methyl for-
Solar System Exploration Division, Code mate, ethyl formate, ketene, formaldehyde, acet-
691, Astrochemistry Laboratory, NASA Goddard aldehyde, formic and acetic acid, glycolaldehyde,
Space Flight Center, Greenbelt, MD, USA ethylene glycol, several nitriles, and possibly gly-
cine. Some of these organics (e.g., the ethers)
could also be formed in the hot gas from those
Keywords intermediaries formed on and sublimated from
the icy mantles on the dust.
Gas-grain chemistry; Ice evaporation; Interstellar Hot cores eventually evolve into ultracompact
molecules; Protostar H II regions as the UV radiation from the massive
central star breaks out into the surrounding envi-
ronment. In the case of low-mass protostars, the
Synonyms hot corino phase is probably terminated by dissi-
pation of the protostellar envelope by accretion
Hot molecular core onto the circumstellar disk and by powerful
outflows.
Definition
See Also
Hot cores are transient regions surrounding mas-
sive ▶ protostars very early in their evolution. ▶ Gas-Grain Chemistry
Similar regions are identified around low-mass ▶ HII Region
protostars and are called hot corinos. Hot cores ▶ Hot Corino
were originally identified as high-temperature ▶ Interstellar Chemical Processes
regions near massive ▶ protostars which showed ▶ Interstellar Dust
high abundances of ammonia. ▶ Interstellar Ices
1120 Hot Corino
▶ Molecular Cloud References and Further Reading

▶ Protostars Cazaux S, Tielens AGGM, Ceccarelli C, Castets A,
Wakelam V, Caux E, Parise B, Teyssier D (2003)
The hot core around the low-mass protostar IRAS
References and Further Reading 16293-2422: Scoundrels Rule! Astrophys J 593:
L51–L55
van Dishoeck EF (2008) Organic matter in space – an
overview. In: Kwok S, Sandford S (eds) Organic mat-
ter in space. Cambridge University Press, Cambridge,
pp 3–16 Hot Jupiters

1
Hot Corino Cambridge, MA, USA
2
Steven B. Charnley Manoa, Honolulu, Hawaii, HI, USA
Space Flight Center, Greenbelt, MD, USA Definition
Hot Jupiter is a designation that is often assigned

Definition to gas giant planets in short-period orbits around
their parent stars. The characteristics of Hot Jupi-
The name hot corino has been given to the imme- ters are not well defined, but the orbital period
diate environment of a low-mass protostar in must be short enough (say less than 10 or 20 days)
which the gas and dust have been heated by the for the planet to be truly “hot,” and the mass must
protostellar radiation and shock waves. Hot be large enough (say more than about one third of
corinos are characterized by large abundances a Jupiter mass) for the planet to be a gas giant. It is
of complex molecules and high ▶ deuterium commonly accepted that Hot Jupiters originally
fractionation ratios, both indicative of sublima- form much farther from their host stars and
tion of ▶ interstellar ices. migrate inwards either by interaction with the
circumstellar disk from which they formed or by
gravitational interactions with other planets in the
History system. For reasons that are not fully understood,
there is a pileup in the number of Hot Jupiters
Hot corinos were so named by Cazaux with orbital periods near 3 days around solar-type
et al. (2003) because of the similarity with the stars. The first Hot Jupiter, which is also the first
hot cores found around high-mass protostars. detected extrasolar planet around Sun-like stars,
was found in 1995, orbiting the star 51 Peg
(Mayor and Queloz 1995).
See Also
▶ Deuterium See Also

▶ Hot Core
▶ Interstellar Ices ▶ Exoplanets, Discovery
▶ Isotopic Fractionation (Interstellar Medium) ▶ Gas Giant Planet
▶ Molecules in Space ▶ Planetary Migration
▶ Protostars ▶ Radial-Velocity Planets
Hot Spring Microbiology 1121

Hot Spring Microbiology
Mayor M, Queloz D (1995) A Jupiter-mass companion to
a solar-type star. Nature 378:355–359
Felipe Gomez
Centro de Astrobiologı́a (CSIC/INTA), Instituto
Hot Molecular Core
▶ Hot Core Definition
Hot spring microbiology refers to the study of

microorganisms associated with hot springs. Hot
Hot Neptunes springs are characterized by hydrothermal sys-
tems that constantly release geothermal gases
David W. Latham1 and Nader Haghighipour2 and fluids from the subsurface due to water H
1
Harvard-Smithsonian Center for Astrophysics, sources in contact with shallow magma forma-
Cambridge, MA, USA tions. Many hot springs have temperatures at or
2
Institute for Astronomy, University of Hawaii- near boiling point; thus, microorganisms from
Manoa, Honolulu, Hawaii, HI, USA these environments are hyperthermophilic or
thermophilic and phylogenetically very diverse,
including ▶ archaea, bacteria, and viruses. Water
Definition chemistry and pH can vary dramatically in these
environments. ▶ Methanogens that belong to the
Hot Neptune is a designation that is often Archaea group and grow at temperature between
assigned to planets with masses or radii similar 80 and 90 C are present in these systems.
to those of Neptune in short-period orbits around Cyanobacteria are often the dominant or sole
their parent stars. The characteristics of Hot Nep- photosynthetic organisms present in hot springs.
tunes are not well defined, but the orbital period Species of Thermotoga capable of growing at
must be short enough (say less than 10 or 20 days) 90 C have been isolated in hot springs and
for the planet to be truly “hot,” and the mass must marine hydrothermal vents. Thermophilic
be substantially smaller than that of Saturn (say Crenarchaeota have been isolated from hot,
less than a tenth of a Jupiter mass). It is com- sulfur-rich solfataras, and some methanogenic
monly accepted that Hot Neptunes originally archaea have also been identified in these envi-
form much farther from their host stars and then ronments. Viruses are represented by extreme
migrate inward either by interaction with the thermophiles, bacteriophages, living up to 85–-
circumstellar disk from which they formed or by 90 C. These microorganisms play important
gravitational interactions with other planets in the roles in the carbon, sulfur, and iron cycles in hot
system. spring ecosystems.
See Also
See Also
▶ Archaea
▶ Exoplanets, Discovery ▶ Crenarchaeota
▶ Gas Giant Planet ▶ Extreme Environment
▶ Planetary Migration ▶ Hot Vent Microbiology
▶ Radial-Velocity Planets ▶ Hyperthermophile
1122 Hot Vent Microbiology
▶ Iron Cycle astrobiological connotations because they are

▶ Methanogens considered interesting models for the origin
▶ Sulfur Cycle of life on Earth as well as potential habitats on
▶ Thermophile other planetary bodies such as Jupiter’s moon,
Europa.
See Also
Hot Vent Microbiology
▶ Archaea
Felipe Gomez
▶ Bacteria
Centro de Astrobiologı́a (CSIC/INTA), Instituto
▶ Barophile
▶ Hyperthermophile
▶ Methanogens
Synonyms
▶ Nitrification
▶ Sulfur Cycle
Hydrothermal vent microbiology
▶ Thermophile
Definition
Hotspot
Hot vent microbiology refers to the microbial
diversity associated with underwater hot springs
▶ Mantle Plume, Planetary
known as hydrothermal vents. Chemical analysis
of hydrothermal fluids shows large amounts
of reduced inorganic materials, including
H2S, Mn2+, H2, NH4, and CO. The system is
based on chemolithoautotrophy (nitrifying Hoyle, Fred
metal- and sulfur-oxidizing microorganisms and
▶ methanogens). Hydrothermal vents combine Fernando B. Figueiredo
several extreme conditions including tempera- CITEUC, University of Coimbra, Coimbra,
ture (up to 380 C), pressure (20 MPa at Portugal
2,000 m), and pH (plumes of pH 2.5), in the
absence of sunlight, all of which determine that
primary producers have to use chemical energy History
for CO2 fixation. The temperature gradient from
the surrounding water that is heated up to Fred Hoyle (1915–2001) besides a scientist was
300–400 C by magma that pours out through also an English national figure, born in June
cracks in the lithosphere to seawater, which 24, 1915, at Gilstead, Bingley West Yorkshire,
remains around 2 C, creates a very unstable England. He graduated at Cambridge Univer-
environment. Chimney growth from mineral pre- sity, where he enrolled in 1933. Despite having
cipitation when hydrothermal fluids come into arrived at Cambridge’s Pembroke College with
contact with cold seawater (black smokers) serious gaps in his mathematical background
serves as a substrate for associated microorgan- training, a fact he quickly surpasses, Fred
isms that can metabolize sulfur and metal sulfide Hoyle would prove to be a brilliant student win-
compounds. The seafloor vents have important ning the Mathematical Tripos in 1936 and being
Hoyle, Fred 1123
awarded with the Mayhew Prize as the best stu- theory that other rare elements could be
dent in applied mathematics. During the Second explained by supernovae.
World War he worked for the Admiralty, Although ironically the term “big bang” was
returning to Cambridge in 1945 as lecturer in coined by Hoyle himself in a BBC’s radio show,
mathematics (1945–1956). Between 1957 and the truth is that Hoyle did not accept the theory of
1972 Fred Hoyle will occupy the Plumian a primordial explosion as being at the origin of
chair, one of the major Professorships in Astron- the universe and proposes a model alternative, the
omy and Experimental Philosophy at Steady State Theory. Hoyle’s name is intimal
Cambridge’s University, and became the foun- related to the concept of continuous creation of
der director of the Institute of Theoretical matter in the universe and to the field equations
Astronomy (1967–1972). that achieved this result. In an attempt to adapt his
Throughout his very active and long scientific primitive ideas to the measurement results on the
live Hoyle will win many awards. He was elected background radiation, which over the years were
Fellow of the Royal Society of London in 1957 being made, he will publish, coauthored with
and foreign associate of the US National Acad- Burbidge and Narlikar, a book, A Different
emy of Sciences, and he was president of the Approach to Cosmology (Cambridge University H
Royal Astronomical Society between 1971 and Press, 2000), presented an alternative to the Big
1973, which had awarded him its gold medal in Bang, by employing an oscillating and expanding
1968. In 1974 he awarded the Royal Medal of the steady-state universe.
Royal Society and in 1997 the Crafoord Prize In 1962 he started a famous collaboration
from the Swedish Academy of Sciences. In with Nalin C. Wickramasinghe on the carbona-
1977 he won the Klumpke-Roberts Prize from ceous nature of interstellar dust and interstellar
the Pacific Astronomical Society, and in 1994, biology. The developments achieved by both
along with Martin Schwarzschild, he would win will lead them to postulate the organic theory
the Balzan Prize. Hoyle was one of the main of cosmic dust, which states among other things
responsible in the planning and construction of that the seeds of life (i.e., the basic chemicals
the Anglo-Australian Telescope when he chaired of DNA replication), including disease viruses,
the Science Research Council’s advisory com- periodically fall from space transported by
mittee. Queen Elizabeth II made him knight in meteorites. To Hoyle life on Earth couldn’t
1972. have arisen by chance in a primordial soup;
In astronomy Hoyle’s works spans from astro- he favored and popularized a view called
physics and particle physics to cosmology. Fred panspermia, the notion that life originated in
Hoyle was the first, in 1946, to establish in solid space and was driven to Earth by electromag-
bases the concept of stars’ nucleosynthesis. In netic radiation pressure.
1957, in a joint article with Margaret Burbidge, Most of his controversial ideas were exposed
Geoff Burbidge, and Willy Fowler, Hoyle will in a lot of his popular non-fiction and science
publish in the Reviews of Modern Physics the fiction books, including Frontiers of Astronomy
famous B2FH paper (the name comes from the (1955), Men and Materialism (1956), Star For-
last initials of the authors). In this work entitled mation (1963), Galaxies, Nuclei and Quasars
Synthesis of the Elements in Stars, the authors (1965), The Relation of Physics and Cosmology
give an explication for the existence in the uni- (1973), Ten Faces of the Universe (1977), and On
verse of elements heavier than helium, showing Stonehenge (1977). In 1994 he published his
that all of the elements from carbon to uranium autobiography, Home Is Where the Wind Blows.
could be produced by nuclear processes in stars, Besides books Fred Hoyle also wrote some plays,
starting with the hydrogen and helium. Hoyle television stories and A for Andromeda serial,
showed that the new young stars are already and an opera, Alchemy Of Love, or the Daemon
formed with these heavy elements, which were Servant’s Retribution, which premiered in New
created in older ones. He also established the York in 1969.
1124 H-Phosphonate
See Also Unlike gas chromatography, the composition of

the mobile phase is often varied with time while
▶ DNA the column temperature is held constant.
▶ Interstellar Dust Reverse-phase HPLC uses a hydrophilic mobile
▶ Panspermia and hydrophobic stationary phase, the opposite of
normal-phase HPLC. HPLC is suited for cou-
pling with different detectors, particularly in
so-called hyphenated techniques, such as HPLC
Burbidge G (2003) Sir Fred Hoyle. Biogr Mem Fellows R coupled to mass spectrometry (LC-MS).
Soc 49:213–247
Burbidge G, Burbidge M (2002) Sir Fred Hoyle
1915–2001. Observatory 122:133–136
Clayton DD (2007) Hoyle, Fred. In: Thomas H et al (eds) See Also
The biographical encyclopedia of astronomers.
Springer, New York, pp 532–533
▶ Chromatography
Lynden‐Bell D (2001) Sir Fred Hoyle (1915–2001).
Observatory 121:405–408 ▶ Ion-Exchange Chromatography
H-Phosphonate
HR 8799: The First Directly Imaged
▶ Phosphite Multi-planet System
Marc Kuchner1 and Nader Haghighipour2

1
NASA Goddard Space Flight Center,
HPLC Exoplanets and Stellar Astrophysics Laboratory,
Greenbelt, MD, USA
2
Jason P. Dworkin Institute for Astronomy, University of Hawaii-
NASA Goddard Space Flight Center, Manoa, Honolulu, Hawaii, HI, USA
Astrochemistry Laboratory, Greenbelt, MD, USA
Definition
Synonyms
HR 8799b, c, d, and e is a system of four planets
High-performance liquid chromatography; High- or possibly ▶ brown dwarfs orbiting HR 8799, a
pressure liquid chromatography; LC; UHPLC young star of about 1.5 solar masses, located
129 light years away in the constellation Pegasus.
The planets were discovered by Christian Marois
Definition and his colleagues using adaptive optics systems
on the Gemini and Keck telescopes in 2008 and
HPLC is an efficient method of liquid–solid col- 2010. The objects b, c, d, and e orbit at distances
umn chromatography. Using a stationary phase roughly 68, 38, 24, and 14.5 AU, respectively,
composed of small (3–10 mm) beads increases from the star (Fig. 1).
separation efficiency but requires pumps to The system of HR 8799 could represent the first
force the mobile phase through the column at multi-planet system discovered via direct imaging.
pressures >300 bar. Ultra-HPLC (UHPLC) uses Mass estimates for the objects based on their lumi-
<2 mm particles and pressures >1,000 bar. nosities and on the age of the star are roughly 7, 10,
HST 1125
HR 8799: The First

Directly Imaged Multi-
planet System,
Fig. 1 Images of the four
planets orbiting star
HR 8799
10, and 10 Jupiter masses for b, c, d, and e, respec-

tively. These values could be roughly 50 % higher, HST
which would make some of them ▶ brown dwarfs,
not planets. However, dynamical arguments sug- Michel Viso
CNES/DSP/SME, Vétérinaire/DVM, Astro/ H
gest that the actual masses are lower than these
numbers, supporting the interpretation that they Exobiology, Paris Cedex 1, France
are all of planetary nature.
HR 8799 also hosts a ▶ debris disk, first dis-
covered by the IRAS satellite in 1986. The most Keywords
recent analysis of the observational data from the
Herschel telescope indicates a planetesimal belt Exoplanet; Planetary disks; Star formation
extending from 100 to 310 astronomical units
(AU) and an external halo extending to approxi-
mately 2000 AU. Synonyms
Hubble Space Telescope

See Also
▶ Adaptive Optics
▶ Brown Dwarf Definition
▶ Direct-Imaging, Planets
▶ Exoplanets, Discovery The Hubble Space Telescope (HST) is a
▶ Fomalhaut b NASA space observatory (Fig. 1) with a 2.5-m-
diameter primary mirror operating in ultraviolet,
visible, and infrared light. Launched in 1990, the
References and Further Reading HST was serviced five times by the space shuttle
and will be operated up to April 2016.
Marois C et al (2008) Direct imaging of multiple planets
The 11,000-kg spacecraft is placed in low
orbiting the star HR 8799. Science 322:1348–1352
Marois C et al (2010) Imaging of a fourth planet orbiting Earth orbit at an altitude of about 575 km. The
HR 8799. Nature 468:1080–1083 data archiving and the scientific operations are
managed by the Space Telescope Science
Institute in the name of the scientific consortium
consisting of numerous universities and institu-
HR Diagram tions. The ▶ European Space Agency contrib-
utes through various participation up to 15 % of
▶ Hertzsprung-Russell Diagram the total cost.
1126 HST
HST, Fig. 1 The Hubble

Space Telescope imaged by
the shuttle during the last
servicing mission in 2009
(Credit NASA)
History Since then and with the following four main-

tenance flights, the HST has delivered unprece-
From 1970 to 1978 US astronomers and NASA dented results in all fields of astronomy. The
deployed an intense lobbying effort toward US overall US expenditure for the HST is estimated
congressmen to have the Large Space Telescope at between $4.5 and $6 billion, with Europe’s
funded. The decision was taken after the decision financial contribution at about €600 million.
of the European Space Agency to support this
effort. Originally slated to be launched in 1983,
technical delays made it almost ready only by Overview
1986. The Challenger accident then grounded
the Space Shuttle fleet for almost 3 years. The The Hubble Space Telescope was sent in orbit on
telescope had to be kept in a clean room, powered April 24, 1990, by the Space Shuttle Discovery.
up, and purged with nitrogen. These operations This 11,000-kg telescope was serviced five times
were costing about $6 million per month, pushing by the space shuttle on its working orbit at an
the overall costs of the project up to $2.5 billion altitude of about 575 km. The telescope was built
by the time of the launch. During this time period, by NASA with a significant contribution from
the engineers performed extensive tests, swapped ESA. It is operated by NASA and the data are
out a possibly failure-prone battery, and made archived at the Space Telescope Science Insti-
several technical improvements. The ground soft- tute in Baltimore, USA. The Hubble’s spectral
ware controlling Hubble was just ready by the range extends from the ultraviolet, through the
1990 launch. visible, and into the near-infrared. Hubble’s pri-
Once in space, the first images were not as sharp mary mirror is 2.4 m in diameter, which is not
as expected and demonstrated a default in the large by ground-based standards. The instru-
primary mirror curvature. This flaw was mainly ments as well as the avionics and hardware
impairing the observation of faint objects and the were upgraded and exchanged during the five
cosmology program. The origin of the flaw was successive servicing missions. Now, and up to
clearly identified and a corrective instrument was the end of the mission, the HST is equipped
designed. Awaiting the repair by the shuttle mis- with five instruments. The Advanced Camera
sion in December 1993, HST performed many for Surveys (ACS) was installed in 2002,
valuable observations for astronomers. improving the field of view and the light
HST 1127
sensitivity. It is working with the newly installed

Wide Field Camera (WFC3). The Cosmic Ori-
gins Spectrograph (COS) instrument is an ultra-
violet spectrograph optimized for observing
faint point sources with moderate spectral reso-
lution. The Near-Infrared Camera and Multi-
object Spectrometer (NICMOS) is the infrared
instrument able to see through interstellar gas
and dust. Finally, the Space Telescope Imaging
Spectrograph (STIS) separates light into compo-
nent wavelengths, acting much like a prism. All
these instruments are placed at the focal plane of
the telescope.
Key Research Findings HST, Fig. 2 This natural-color image of Jupiter displays at H
bottom right debris from a comet or asteroid that plunged
into Jupiter’s atmosphere and disintegrated (Credit: NASA,
The HST was built and designed to image far, ESA, Michael Wong [Space Telescope Science Institute,
faint galaxies. This fossil light reveals how the Baltimore, MD], H. B. Hammel [Space Science Institute,
universe looked in the remote past and how it Boulder, CO], and the Jupiter Impact Team)
may have evolved with time. The Hubble Deep
Fields gave a clear glance back to the time when
galaxies were forming. Deep field observations star (▶ HD 209458b) located 150 light-years
require long-lasting pointing accuracy to a away in the constellation Pegasus.
selected region of the sky. Deeper observation Hubble made also unique, repeatable observa-
requires longer exposure time, while the fainter tions of the surfaces of the planets and objects of
objects become visible on the images. The first our Solar System: imaging the dust storms on
deep fields gave new insight about the early Mars, preparing for the flyby of asteroids like
Universe, revolutionizing modern astronomy. Vesta by the Dawn mission, or recording “live”
The HST has harvested outstanding results in the diving of the comet Shoemaker-Levy into the
many fields of astronomy, like the calculation of atmosphere of Jupiter (see Fig. 2).
the Hubble constant, the evaluation of the age of
the Universe, and the astrophysics of objects
such as quasars and supermassive black holes Future Directions
in the centers of galaxies. The systematic sur-
veys were also used to detect gravitational The James Webb Space Telescope (▶ JWST), to
lenses and map the distribution of dark matter be placed in an orbit at the Lagrangian point L2, is
in some portions of the Universe. For astrobiol- planned to replace the Hubble Space Telescope.
ogy, the Hubble Space Telescope has contrib-
uted to the imaging of the clouds of gas and dust
where new stars are forming. Data collected See Also
from the Orion Nebula are giving new insight
into these processes. The detected dust disks ▶ Comet Shoemaker-Levy 9
around newborn stars could be the beginning ▶ European Space Agency
of planetary systems. Hubble performed the ▶ HD 209458b
first detection of an atmosphere around an ▶ JWST
extrasolar giant planet, which orbits a Sunlike ▶ NASA
1128 Hubble Space Telescope
Hubble Space Telescope Huronian Glaciation
▶ HST Andrey Bekker

Department of Geology and Geophysics,
Yale University, New Haven, CT, USA
Hubble, Edwin
Keywords
CITEUC, University of Coimbra, Coimbra, Paleoproterozoic; Ice ages; Snowball earth; Rise
Portugal of atmospheric oxygen
History Synonyms
Edwin Powell Hubble (1889–1953) was an Paleoproterozoic ice ages; Paleoproterozoic

American astronomer known by his works on snowball earth
galactic and extragalactic astronomy. His works
provided the first persuasive observational evi-
dence that the Universe is expanding. Definition
In 1929, with data collected from the best
telescope of that time, the 100-in. telescope of The Huronian glaciation is the oldest series of
the Mount Wilson Observatory, he showed that protracted climatic refrigeration events that
galaxies are moving away from Earth with a extensively affected Earth. It occurred between
speed proportional to their distance according 2.45 and 2.22 Ga in association with the rise of
to the law, v = H0 d, where v is the recession atmospheric oxygen. Three glaciations of that
velocity of a galaxy at a distance d and H0 is the series, the classical Huronian ice ages, are brack-
value for the Universe expansion rate at the eted in time between ~2.45 and 2.32 Ga; the
current epoch. Hubble’s initial value for H0 fourth event, recognized so far only in South
(now called the Hubble constant) was calculated Africa, is ~2.22 Ga in age. During these events,
to be approximately 500 km s1 Mpc1. glaciers covered continents, extended to low lat-
The Hubble Space Telescope, a satellite itudes, and reached to sea level. The ice ages were
observatory orbiting Earth, launched by followed by a protracted time interval with green-
NASA’s Space Shuttle mission STS-31 on house (warm and humid) conditions. The name is
April 24, 1990, is continuing the work begun derived from the Huronian Supergroup exposed
by Hubble himself to study and map our Uni- on the north shore of Lake Huron in Ontario,
verse and helping to determine the Hubble con- Canada, between Sault Ste. Marie, Sudbury, and
stant to an accuracy of
10 %; the actual H0 Cobalt.
value is 72
8 km s1 Mpc1.
History
The Huronian glacial deposits were first recog-

Hungarian-Made Automatic nized by Coleman (1907) in the northeastern part
Telescope Network of the Huronian Basin in Ontario, Canada; just a
year after a poorly sorted conglomerate with
▶ HATNet scattered pebbles and cobbles, some striated and
Huronian Glaciation 1129
Huronian Glaciation,
Fig. 1 Location of
Paleoproterozoic glacial
deposits in North America
and Fennoscandia
(Modified after Young
2004). Abbreviations
represent stratigraphic units
which contain glacial
deposits. S Snowy Pass
Supergroup, Medicine Bow
Mountains, Wyoming, Ch
Chocolay Group,
Michigan, Hn Huronian
Supergroup, Ontario,
C Chibougamau
Formation, Quebec, Hw
Hurwitz Group, Nunavut,
K Karelian Supergroup,
Fennoscandia
H
faceted, was described in the Northern Cape above thick banded iron formation associated
Province of South Africa by Rogers (1906), who with mafic volcanics (Mikhalsky et al. 2006;
interpreted it to be glacial in origin. Almost Phillips et al. 2006), offering a comparison with
40 years later, Pettijohn (1943) described tillite the Meteorite Bore Member of the Turee Creek
in a correlative to the Huronian Supergroup suc- Group in the Hamersley Province of Western
cession of Michigan, USA. Young (1970) Australia (see below). All other reported cases
inferred glacial influence on Paleoproterozoic of Paleoproterozoic glacial deposits were either
successions in Wyoming, Michigan, Quebec, not confirmed by subsequent sedimentologic
and Nunavut. Shortly thereafter, Paleopro- studies or turned out to be significantly younger
terozoic glacial deposits were for the first time and not correlative with the Huronian glaciation.
recognized in Western Australia (Trendall 1976)
and described in more details in South Africa
(Visser 1971). Although the presence of Overview
Paleoproterozoic glacial deposits in
Fennoscandia was originally discussed by Eskola Paleoproterozoic glacial deposits are now con-
(1919), they were not described and documented firmed in a number of basins in North America
in detail until recently (Marmo and Ojakangas (Fig. 1), Western Australia (Meteorite Bore
1984; Strand and Laajoki 1993). In Antarctica, Member of the Turee Creek Group), South Africa
the 2.45–2.5 Ga Ruker Series contains diamictite (Pretoria and Postmasburg Groups), and
1130 Huronian Glaciation
Huronian Glaciation, Fig. 2 Sedimentary features of 2.8 cm in diameter. (d) Red sandstone sandwiched
Paleoproterozoic successions indicating glacial influence between glacial diamictites of the Gowganda Formation,
and postglacial warm and humid climate associated with north of Elliott Lake, person for scale. (e) Molds of gyp-
oxygenated atmosphere. (a) Dropstone in finely laminated sum in red-colored siltstone from postglacial succession in
mudstone of the Pecors Formation, Huronian Supergroup, Michigan (HW 480; Kona Dolomite); coin is 2.8 cm in
north of Quirke Lake; hammer for scale is 40 cm long. (b) diameter for scale. (f) Cross-bedded red-colored
Large dropstone in laminated matrix of the Gowganda mudstone-siltstone in the upper part of the Gowganda
Formation in Cobalt area; hammer is 70 cm long. (c) Formation deposited in oxygenated deltaic setting, Cobalt
Dropstone of granite in finely laminated matrix of the area; hammer for scale is 70 cm long
Gowganda Formation, north of Elliott Lake; coin is
Fennoscandia (Sarioli informal group in Finland contains three glacial units (Fig. 3), whereas other
and in Karelia and Kola Peninsula, Russia). Evi- successions contain only one or two, making
dence for glaciation is plentiful in these succes- intra- and intercontinental lithostratigraphic cor-
sions and includes varves with dropstones; relations difficult. It seems likely that four glacial
striated, flat-iron-shaped, and faceted pebbles; events occurred in the Paleoproterozoic, but their
striations in the underlying basement; exotic regional or global extent is not well constrained.
stones in diamictite (Fig. 2); and extensive Paleomagnetic data are as yet scarce; however,
development of diamictite not controlled by they suggest low-latitude position of glaciated
synsedimentary faults. The Huronian Supergroup landmasses (e.g., Evans et al. 1997), whereas
Huronian Glaciation
Huronian Glaciation, Fig. 3 Age-calibrated correlation of Paleoproterozoic strata are in bold; also shown are age constraints from previous studies (see Rasmussen
from North America and southern Africa (updated from Rasmussen et al. 2013 with U/ et al. 2013 for references) and the age of the Copper Cliff rhyolite at the base of the
Pb baddeleyite age from Gumsley et al. 2015; multiple S isotope data and detrital Huronian Supergroup (in italics; Ketchum et al. 2013). Values in blue adjacent to
pyrite and uraninite description from Johnson et al. 2013, 2014; and evidence for marine carbonate units represent their carbon isotope compositions. MIF and MDF are
redbeds from Schröder et al. 2011). Dates from Rasmussen et al. (2013) for tuff beds mass-independent and mass-dependent fractionations of sulfur isotopes, respectively
1131
H
Huronian Glaciation, Fig. 4 Secular carbon isotope ca. 2.07 Ga indicates that sulfate evaporites again became
variations in seawater and redox indicators for the oxida- rare in the Paleoproterozoic and Mesoproterozoic record
tion state of the early Paleoproterozoic atmosphere-ocean after that time. Deposition of iron formations and Mn-rich
system (Modified from Bekker and Holland 2012). Four deposits indicates anoxic conditions in deep waters. While
blue vertical bars mark Paleoproterozoic glacial events; deposition of iron formations does not necessarily require
the dashed secular carbon isotope curve between 2.5 and atmospheric oxygen and can be mediated by anoxygenic
2.22 Ga emphasizes the uncertainty in this part of the photosynthetic bacteria, Mn oxidation requires significant
curve, the dashed bar for marine sulfate evaporites after levels of atmospheric oxygen
sedimentologic data indicates that ice extended to else in the world. The Makganyene Diamictite
sea level in these regions (e.g., Young 2004). of the Postmasburg Group of the Griqualand
Large igneous provinces were extensively West basin (paleomagnetically pinned to low
emplaced before (2.5–2.45 Ga) and after paleolatitudes; Evans et al. 1997) was recently
(ca. 2.22 Ga) the Huronian glaciation, but with geochronologically bracketed between ~2.46 and
the exception of the ca. 2.32 Ga superplume 2.43 Ga in age (see Fig. 3; Gumsley et al. 2015),
event, the 2.45–2.22 Ga time interval is largely potentially providing the best age constraint for
devoid of magmatic activity (cf. Partin the beginning of the GOE. Correlation of the
et al. 2014). Tuff beds dated in the Huronian glacial diamictite of the Meteorite Bore Member
Supergroup, Canada, and the Pretoria Group, of the Turee Creek Group with the Huronian
South Africa (Rasmussen et al. 2013), have glacials remains uncertain; considering that the
finally bracketed the age of the Paleoproterozoic retro-arc setting of the Turee Creek Group
ice ages (Fig. 3). Three Huronian glaciations are evolved from a conformably underlying back-
all between ca. 2.45 and 2.32 Ga in age, whereas arc basin containing 2.5–2.45 Ga banded iron
the youngest glaciation recorded by the upper formations (e.g., Krapež 1996), the most parsi-
Pretoria Group is ca. 2.22 Ga in age. These data monious interpretation is that it represents the
allow two tantalizing implications: (1) The oldest Huronian ice age.
Huronian ice ages lasted long; (2) there was a In contrast to Neoproterozoic glacially
glacial event at ca. 2.22 Ga in South Africa, influenced successions, carbonates with negative
which has not yet been recognized anywhere carbon isotope values directly overlying glacial
Huronian Glaciation 1133
Huronian δ13C, ‰ V-PDB

Supergroup,
ON, Canada 2217.5 ± 1.6 Ma
Nipissing
Bar
intrusions
River Fm.
Gordon
Lake Fm. +5.0 to +8.2 ‰ 2308 ± 8 Ma sulfate evaporites
Cobalt Gr.
Lorrain Mature Quartzites / Red Beds

Fm.
oxidized paleosols
Gowganda
Fm. 3rd glaciation
LEGEND:
diabase dikes and sills
Serpent
Quirke Lake
Fm. carbonate
−4.0 to −0.8 ‰
Gr.
Espanola mature quatzite with red beds

Fm. cap carbonate
diamictite
Bruce Fm. 2nd glaciation H
shale and siltstone
Mississagi
Hough Lake
mafic volcanics
Fm.
sandstone
Gr.
Pecors Fm. conglomerate, sandstone,

Ramsay and siltstone
Lake Fm. 1st glaciation
basement
McKim
Elliot Lake
Fm. unconformity
Gr.
Matinenda detrital pyrite and uraninite

Fm.
Reduced paleosols
2452.5 ± 6.2 Thessalon Fm. 2.43-2.5 Ga Tectonomagmatic event
Ma
Late detrital pyrite and uraninite
Archean Livingstone Creek Fm.
Huronian Glaciation, Fig. 5 Stratigraphic column of atmospheric redox indicators, age constraints, and envi-
the Huronian Supergroup in Ontario, Canada that contains ronmental and tectonic changes (Modified from Bekker et
three glacial horizons and overlying units indicating al. 2006 and Rasmussen et al. 2013)
extreme degree of weathering. Also shown are
diamictite are present only above the second gla- overlying units contain red beds and sulfate evap-
cial diamictite in the Huronian Supergroup and orites (Figs. 2 and 4). Combined, these and other
correlative glacial horizons elsewhere. There are proxies for atmospheric and ocean redox state
carbonates with negative carbon isotope values indicate a transition from the reduced to an oxy-
immediately underlying the glacial diamictite of genated atmosphere and ocean during the
the Polisarka Sedimentary Formation on the Kola Huronian glaciations (Bekker et al. 2004). It is
Peninsula of Russia, but correlation of this glacial generally accepted that the atmosphere before
diamictite with the Huronian glacials and its age ca. 2.45 Ga was reducing and contained signifi-
is uncertain (Brasier et al. 2013). Sequences cant ( 1,000 ppm) levels of methane, an effec-
immediately underlying Paleoproterozoic glacial tive greenhouse gas. Rise of atmospheric oxygen
diamictites include banded iron formations in in the early Paleoproterozoic decreased methane
Western Australia and South Africa and con- levels in the atmosphere and initiated climatic
glomerates with detrital pyrites and uraninites in cooling leading to glaciation (Bekker et al.
North America and ▶ Fennoscandia, whereas 2005). If this interpretation is correct, the
Huronian glaciation represents a response of Coleman AP (1907) A lower huronian ice age. Am J Sci
Earth system to the rise of atmospheric oxygen 23:187–192
Eskola PE (1919) Hufvuddragen av Onega-Karelens geol-
in association with carbon dioxide replacing ogy: Helsingin Geol. Yhd. Tiedonantoja 1917 u. 1918,
methane as a main greenhouse gas. Lithologies pp 13–18, and Teknikern, vol 29, pp 37–39
overlying Paleoproterozoic glacial deposits sug- Evans DA, Beukes NJ, Kirschvink JL (1997) Low-latitude
gest an extended period of enhanced chemical glaciation in the Paleoproterozoic era. Nature
386:262–266
weathering in the aftermath of the Huronian gla- Gumsley AP, Chamberlain K, Bleeker W, Söderlund U, de
ciations, which could reflect warm and humid Kock MO, Kampmann TC, Larsson E (2015) U-Pb
climate and acidic groundwaters during the TIMS and in-situ SIMS dating of baddeleyite and
Great Oxidation Event and Lomagundi carbon zircon from sub-volcanic sills of the Ongeluk Forma-
tion (Transvaal Supergroup) in the Griqualand West
isotope excursion (Konhauser et al. 2011; Bekker sub-basin, Kaapvaal Craton, with implications for
and Holland 2012) (Fig. 5). Snowball Earth and the Great Oxygenation Event.
AGU-GAC-MAC-CGU Joint Assembly 2015 Pro-
gram with Abstracts.
See Also Johnson JE, Gerpheide A, Lamb MP, Fischer WW (2014)
O2 constraints from Paleoproterozoic detrital pyrite
and uraninite. GSA Bull 126(5/6):813–830
▶ Cap Carbonates Johnson JE, Webb SM, Thomas K, Ono S, Kirschvink JL,
▶ Diamictite/Diamicton Fischer WW (2013) Manganese-oxidizing photosyn-
▶ Glaciation thesis before the rise of cyanobacteria. PNAS 110
(28):11238–11243
▶ Great Oxygenation Event Ketchum KY, Heaman LM, Bennett G, Hughes DJ
▶ Lomagundi Carbon Isotope Excursion (2013) Age, petrogenesis and tectonic setting of the
▶ Oxygenation of the Earth’s Atmosphere Thessalon volcanic rocks, Huronian Supergroup, Can-
▶ Paleosols ada. Precambrian Res 233:144–172
Konhauser KO, Lalonde SV, Planavsky NJ, Pecoits E,
▶ Precambrian Lyons TW, Mojzsis SJ, Rouxel OJ, Barley ME,
▶ Proterozoic Eon Rosı̀ere C, Fralick PW, Kump LR, Bekker A (2011)
▶ Pyrite Aerobic pyrite oxidation and acid rock drainage during
▶ Red Beds the great oxidation event. Nature 478:369–373
Krapež B (1996) Sequence-stratigraphic concepts applied
▶ Sedimentary Rock to the identification of basin-filling rhythms in Precam-
▶ Shale brian successions. Aust J Earth Sci 43:355–380
▶ Snowball Earth Marmo JS, Ojakangas RW (1984) Lower Proterozoic
▶ Stratigraphy glaciogenic deposits, eastern Finland. GSA Bull
95:1055–1062
▶ Transvaal Supergroup, South Africa Mikhalsky EV, Beliatsky BV, Sheraton JW, Roland NW
(2006) Two distinct Precambrian terranes in the South-
ern Prince Charles Mountains, East Antarctica:
References and Further Reading SHRIMP dating and geochemical constraints. Gond-
wana Res 9:291–309
Bekker A, Holland HD (2012) Oxygen overshoot and Partin CA, Bekker A, Sylvester PJ, Wodicka N, Stern RA,
recovery during the early Paleoproterozoic. Earth Chacko T, Heaman LM (2014) Filling in the juvenile
Planet Sci Lett 317–318:295–304 magmatic gap: evidence for uninterrupted Paleopro-
Bekker A, Holland HD, Wang P-L, Rumble D III, Stein terozoic plate tectonics. Earth Planet Sci Lett
HJ, Hannah JL, Coetzee LL, Beukes NJ (2004) Dating 388:123–133
the rise of atmospheric oxygen. Nature 427:117–120 Pettijohn FJ (1943) Basal Huronian conglomerates of
Bekker A, Kaufman AJ, Karhu JA, Eriksson KA Menominee and Calumet Districts, Michigan. J Geol
(2005) Evidence for Paleoproterozoic cap carbonates 51:387–397
in North America. Precambrian Res 137:167–206 Phillips G, Wilson CJL, Campbell IH, Allen CM
Brasier AT, Martin AP, Melezhik VA, Prave AR, Condon (2006) U-Th-Pb detrital zircon geochronology from
DJ, Fallick AE, FAAR-DEEP Scientists (2013) Earth’s the southern Prince Charles Mountains, East
earliest global glaciation? Carbonate geochemistry Antarctica – defining the Archean to Neoproterozoic
and geochronology of the Polisarka Sedimentary For- Ruker Province. Precambrian Res 148:292–306
mation, Kola Peninsula, Russia. Precambrian Res Rasmussen B, Bekker A, Fletcher IR (2013) Correlation of
235:278–294 Paleoproterozoic glaciations based on U-Pb zircon
Huygens 1135
ages for tuff beds in the Transvaal and Huronian with a nucleus, is the formal basis of all life, and
Supergroups. EPSL 382:173–180 he insisted on the fact that materialism is the
Rogers AW (1906) The glacial beds in the Griqua Town
Series. Rep S Afr Assoc Adv Sci 4:261–265 best way to understand life. In the second
Schröder S, Bedorf D, Beukes NJ, Gutzmer J (2011) From text, he strictly rejected the spontaneous gener-
BIF to red beds: Sedimentology and sequence stratig- ation and described the debate on this
raphy of the Paleoproterozoic Koegas Subgroup matter. He suggested distinguishing two terms:
(South Africa). Sedimentary Geology 236:25–44
Strand KO, Laajoki K (1993) Palaeoproterozoic biogenis, when living matter comes from living
glaciomarine sedimentation in an extensional tectonic matter, and abiobenesis, synonymous of spontane-
setting: the Honkala Formation, Finland. Precambrian ous generation, when living matter comes from
Res 64(p):253–271 inert matter.
Trendall AF (1976) Striated and faceted boulders from the
Turee Creek Formation – evidence for a possible
Huronian glaciation on the Australian continent. Geol
Surv West Aust Annu Rep 1975:88–92
Visser JNJ (1971) The deposition of the Griquatown gla- See Also
cial member in the Transvaal Supergroup. Trans Geol
Soc S Afr 74:187–199 ▶ Cellular Theory, History of H
Young GM (1970) An extensive early proterozoic glacia-
tion in North America? Palaeogeogr Palaeoclimatol
Palaeoecol 7:85–101 ▶ Haeckel’s Conception of Origins of Life
Young GM (2004) Earth’s earliest glaciations: tectonic ▶ Protoplasmic Theory of Life
setting and stratigraphic context of Paleoproterozoic
glaciogenic deposits. In: Jenkins GS, McMenamin
MAS, McKay CP, Sohl L (eds) The extreme Protero-
zoic: geology, geochemistry, and climate, AGU geo-
physical monograph series. American Geophysical
Union, Washington, DC, 146, pp 161–181
Huygens
Therese Encrenaz
Huxley’s Conception on Origins of
Life
Stéphane Tirard Definition

des Techniques EA 1161, Faculté des Sciences et Christiaan Huygens (1629–1695) was born in
des Techniques de Nantes, Nantes, France The Hague (Netherlands). He studied law and
mathematics in Leiden and then studied and
built microscopes and telescopes. With his own
History telescope, he discovered ▶ Titan, ▶ Saturn’s
biggest satellite, in 1655. In 1659, he found the
Thomas Huxley (1825–1895) was a British explanation of the changing appearance of Sat-
biologist who endorsed Charles Darwin’s the- urn, due to its rings, which he published in
ory of evolution. As a zoologist, physiologist, Systema Saturnium. In 1656, he built the
and evolutionist, Huxley explained his concep- first pendulum clock with the objective of deter-
tions on nature of living matter in two important mining longitudes at sea. He developed in his
lectures: The Physical Basis of Life (1868) and Traité de la Lumière the first wave theory of
Biogenesis and Abiogenesis (1870). In the first light. His posthumous work, Cosmotheoros,
lecture, he described and analyzed the impor- published in 1698, deals with the nature of the
tance of the protoplasm as the main constituent universe and the habitability of other worlds.
of cells. He wrote that the protoplasm, simple or The probe of the Cassini-Huygens mission,
1136 Huygens Probe
which landed on Titan’s surface on January History

14, 2005, was named after him.
As early as 1982, an ambitious and international
mission to explore the Saturn system was initially
See Also
proposed to ▶ NASA and ESA by a team of
European and US scientists. A probe to specifi-
▶ Saturn
cally study Titan was identified very early in the
▶ Titan
study phase of the mission as a potential contri-
bution of ESA. After a Phase A study, the Titan’s
probe was selected in 1988 by the Science
Huygens Probe Programme Committee of ESA and named “Huy-
gens” at that time. The selection of the ESA
François Raulin Huygens probe investigations, as that of the
Faculté des Sciences et Technologie, Université NASA Cassini orbiter, was done jointly by ESA
Paris Est Créteil et Paris Diderot, LISA – UMR and NASA and announced in the fall of 1990.
CNRS 7583, Creteil, France
Overview
Keywords
The Huygens probe (Lebreton and Matson 2002)
ACP; Atmospheric probe; Cassini-Huygens; was one of the key elements of the Cassini-
DISR; DWE; GC-MS; HASI; SSP; Titan; Titan’s Huygens mission. It was provided by ESA and
atmosphere; Titan’s surface constructed by an industrial consortium, with
Aerospatiale, Cannes, France (currently Thales
Alenia Space), as the prime contractor (Clausen
Synonyms et al. 2002). It was carried by the Cassini space-
craft from its launch, on October 15, 1997, to its
Cassini Titan’s probe; Huygens spacecraft; release, on December 25, 2004. It separated from
Titan’s atmosphere module; Titan’s atmosphere Cassini that day at 2:00 UTC with a relative speed
probe of 0.35 m/s. It entered Titan’s atmosphere on
January 14, 2005, targeted to land on a southern-
latitude site on Titan’s dayside. Protected by its
Definition heat shield (Fig. 1), it experienced a strong
aerobraking, with a velocity decreasing from
The Huygens probe was an atmospheric probe 21,600 km/h during entry to less than
supplied by the European Space Agency (ESA) 1,400 km/h at 180 km altitude and only
and part of the Cassini-Huygens mission to the 290 km/h at 160 km. There the main parachute
▶ Saturn system. The probe was named after the (8.3 m in diameter) opened and then discarded at
Dutch astronomer Christiaan Huygens who about 110 km altitude and replaced by a small one
discovered ▶ Titan, the largest satellite of (3 m in diameter). This last parachute allowed the
Saturn, in 1655. It was designed to explore the probe to reach Titan’s surface before the Cassini
environment of Titan, mainly its atmosphere, spacecraft passed below the horizon of the
which makes Titan so unique. It was carried by ▶ landing site. However, it was efficient enough
the ▶ Cassini spacecraft. It entered Titan’s atmo- to allow a low-velocity landing (5 m/s). The
sphere on January 14, 2005, and studied Titan for descent time in the atmosphere was close to 2 h
several hours during its descent and after its and 30 min. The probe continued operating after
touchdown with the six instruments of its scien- landing on the surface for about 3 h. The first 1 h
tific payload (Table 1). and 12 min of surface data was collected by
Huygens Probe 1137
Huygens Probe, Table 1 The Huygens probe’s scientific investigations and their expected astrobiological return
Huygens instruments and interdisciplinary programs P.I., team leader or IDS Astrobiological return
Scientific instruments
Gas chromatograph-mass spectrometer (GC-MS) H. Niemann USA +++
Aerosol collector and pyrolyzer (ACP) G. Israēl France +++
Huygens atmospheric structure instrument (HASI) M. Fulchignoni Italy ++
Descent imager and spectral radiometer (DISR) M. Tomasko USA +++
Doppler wind experiment (DWE) M. Bird Germany +
Surface science package (SSP) J. Zarnecki U.K. +++
Interdisciplinary scientists
Aeronomy D. Gautier France ++
Atmosphere/surface interactions J.I. Lunine USA +++
Chemistry and exobiology F. Raulin France +++
Huygens Probe, H
Fig. 1 The back cover of
the probe, with its gold-
colored blanket, is attached
to the Huygens probe,
already inside its front
shield (covered by heat-
resistant tiles) (Credit: ESA
(ID number:
SEMJRJ2VQUD))
Cassini, before it went out of the field of view of Titan’s atmosphere, mainly from 150 km alti-
Huygens (Fig. 2). The total mass of the probe was tude, down to the surface, and to, eventually,
about 350 kg including the probe structure, sup- carry out some surface measurements, although
port equipments, and 48 kg of scientific instru- it was not conceived to survive the surface
ments. The probe was developed to explore impact. Huygens included six scientific
1138 Huygens Probe
Huygens Probe, Fig. 2 Artist’s picture of the descent and landing of the Huygens probe on Titan on January 14, 2005
(Credit: ESA-D. Ducros (ID number: SEMF4XULWFE))
experiments (Table 1) to study the physical and polarized transmitting antennas. The data were
chemical properties of the atmosphere and sur- transmitted from Huygens to Cassini on two
face of Titan. nearly redundant channels.
Huygens included a CD-ROM with about The ▶ gas chromatograph-mass spectrometer
100,000 messages collected by ESA between (GC-MS) (Niemann et al. 2002) included three
December 1996 and February 1997. Those mes- GC columns, using dihydrogen as carrier gas,
sages included texts, drawings, audio messages, linked to a quadrupole mass spectrometer and a
and signatures from tens of thousands of citizens gas sampling system. The GC achieved the sepa-
from the planet Earth (Fig. 3). ration of the different constituents of complex
mixtures (atmospheric gases and heated or pyro-
lyzed aerosols from aerosol collector and pyro-
Basic Methodology lyzer [ACP]). The MS allowed their detection
and identification, with a mass range extending
The energy on Huygens was delivered by five from 2 to 141 Da. The instrument was equipped
LiSO2 batteries, capable of delivering more than with five ion sources feeding the mass analyzer
2 KWh. The data (housekeeping and scientific) sequentially. It also included an enrichment cell
were processed by a Command and Data Man- to concentrate trace gases and increase the limit
agement Unit, which included a 16-bit micropro- of detection.
cessor (running at 10 MHz). They were then sent The ACP (Israel et al. 2002) was devoted
to the Cassini orbiter in its solid-state memory to the in situ study of Titan’s aerosols: their
with the Probe Data Relay Subsystem, which chemical composition and their distribution.
included S-band transmitters and circularly It included a sampling system with a filter to
Huygens Probe 1139
and main composition of the atmosphere, as

well as the distribution and properties of the aero-
sols. It was also able to provide many images and
information on nature of the surface.
The Huygens Atmospheric Structure Instru-
ment (HASI; Fulchignoni et al. 2002) was com-
posed of several sensors to measure physical
properties of the atmosphere: accelerometer,
temperature and pressure sensors, a microphone,
and an electric field sensor. One of the main goals
of the HASI was to measure the vertical temper-
ature/pressure atmospheric profile from the
mid-stratosphere to the surface of Titan.
The Surface Science Package (SSP; Zarnecki
et al. 2002) was also a multisensor experiment to
measure surface properties: temperature, thermal H
conductivity, heat capacity, speed of sound,
dielectric constant, and index of refraction
(if liquid). It included a force transducer to mea-
sure the impact deceleration.
The Doppler Wind Experiment (DWE; Bird
et al. 2002) was designed to determine the direc-
tion and strength of the zonal winds in Titan’s
atmosphere. It was composed of a transmitter on
the Huygens probe and a receiver on the Cassini
orbiter, allowing Doppler tracking of the probe
from the orbiter.
Huygens Probe, Fig. 3 The Huygens CD-ROM: front

cover and CD (Courtesy of NASA/JPL/Caltech/ESA) Key Research Findings
collect atmospheric aerosols and an oven to heat The whole Huygens operations worked perfectly,
the collected particles at different temperatures. from the pre-entry in Titan’s atmosphere to the
Once the collecting phase was over, the filter was landing (Lebreton et al. 2005, 2009; Owen 2005).
retracted inside the probe in the oven. Then the The latter was much smoother than expected, and
oven was isolated and the filter heated at different the probe survived the impact and its instruments
temperatures. The produced gases were then ana- continued working for several hours on the sur-
lyzed by the GC-MS instrument. Two samplings face. Only one problem was experienced: the loss
were carried out in the 130–35 km altitude and of one of the two channels because of a program-
the 25–20 km altitude, respectively. ming error in its receiver. Consequently, no data
The Descent Imager and Spectral Radiometer were collected from this channel, which affected
(DISR; Tomasko et al. 2002), the only optical mainly the DWE experiment, but finally most of
remote sensing experiment aboard Huygens, the data were recovered, thanks to Earth-based
operated in the 0.3–1.7 mm wavelength range. It radio telescopes that were able to receive the
included photometers to look upward and down- weak radio signal from Huygens.
ward, solar visible and infrared spectrometers, The scientific results of the Huygens instru-
and several imagers. Its scientific objectives ments were first published in Nature in 2005 and
were to study the thermal balance, dynamics, then in two special issues of Planetary and Space
1140 Huygens Probe
Huygens Probe, Fig. 4 Channel networks, highlands,

and dark-bright interface on the surface of Titan as seen by
the DISR instrument on Huygens at 6.5 km altitude (Credit:
ESA/NASA/JPL/University of Arizona – PIA07236)
Science (Raulin et al. 2007, 2008). They are also

presented together with the Cassini data in a book
Huygens Probe, Fig. 5 Image of Titan’s surface where
devoted to Titan (Brown et al. 2009). The DISR
the Huygens probe landed taken by the DISR instrument
(Tomasko et al. 2005) provided more than on January 14, 2005. It shows pebble-sized objects, likely
350 images and many spectra of Titan. Its images to be made of water ice (Credits: ESA/NASA/JPL/Uni-
did not show any liquid ▶ hydrocarbon pool on versity of Arizona (PIA07232))
Titan’s surface, but dendritic structures formed
by liquid which flowed in the past. It goes from
brighter highland regions down to dark and flat (Zarnecki et al. 2005) performed physical mea-
lowlands (Fig. 4). They also showed that the surements just above and on Titan’s surface. It
Huygens landing site looked like a dry riverbed evidenced a relatively soft solid surface. The
strewn with pebbles probably made of ▶ water DWE team (Bird et al. 2005) was able to deter-
ice (Fig. 5). The DISR infrared spectra of the mine a high-resolution profile of the winds,
surface suggested a mixture of water ice and including evidence for low-velocity winds
organic compounds, including Titan tholin-like below 5 km altitude, and to confirm the super-
material. The DISR data also showed that haze rotation of Titan. The GC-MS (Niemann
particles in the atmosphere seemed continuously et al. 2005) carried out a direct MS analysis of
distributed from 150 km altitude to the surface, Titan’s atmosphere from 146 km altitude down to
where the ▶ methane relative humidity reaches the surface. It determined the concentration pro-
50 %. The HASI (Fulchignoni et al. 2005) deter- files of several atmospheric gases as a function of
mined the temperature, pressure, and density pro- altitude and measured ▶ isotopic ratios and trace
files of the atmosphere from 1400 km to the species. It detected 40Ar as the main noble gas,
surface (Fig. 6). It detected an ionospheric layer with an abundance more than 100 times higher
in the low atmosphere, between 140 and 40 km, than that of 36Ar. It measured 14N/15N and
12 13
with a potential lightning signal. The SSP C/ C and D/H in the atmosphere and found
Huygens Probe 1141
Huygens Probe, Pressure, mbar Altitude, km

Fig. 6 Vertical
temperature and pressure 1,400
profile (dotted line) of 10−8
Titan’s atmosphere from
HASI data (Credit:
ESA/ASI/UPD/OU/FMI)
Thermosphere 1,000
10−6
10−4
Mesopause
500
10−2
Mesosphere
Stratopause
250 H
100
Stratosphere
Tropopause 50
102
Troposphere
0
50 100 150 200 250
Temperature, K
values close to but smaller than the terrestrial NH2

values, except for D/H. It detected organic species
on the surface, including ethane, cyanogen, and CN
benzene. It also observed the presence of methane HN

degassing on the surface. The ACP (Israel
et al. 2005), using GC-MS as an analytical tool,
was able to collect and pyrolyze atmospheric aero- CN
sols in the low stratosphere and mid-troposphere
regions. It showed that the collected particles
release NH3 and HCN when heated at 600 C,
indicating that they are made of refractory NH
organics, including ▶ carbon (C), ▶ hydrogen Refractory
(H), and ▶ nitrogen (N) atoms (Fig. 7). Small fraction
C,H,N
of condensates
organics
Applications
Huygens Probe, Fig. 7 Elemental composition, struc-
The Huygens probe was the first object launched
ture, and example of the possible molecular composition
by mankind that was able to softly land on of Titan’s aerosols derived from the Cassini-Huygens
a planetary object of the outer solar system. ACP data
1142 Huygens Probe
The Huygens landing site is now named the References and Further Reading
Hubert Curien Memorial Station, in recognition
of one of the founders of ESA. The technical and Bird MK et al (2002) The Huygens Doppler wind
experiment’ titan winds derived from probe radio
scientific achievement of Huygens shows our
frequency measurements. Space Sci Rev 104:
capability to land on faraway worlds. It marks 613–640
the beginning of the systematic exploration of Bird M et al (2005) The vertical profile of winds on Titan.
outer solar system objects. Nature 438:800–802
Brown R, Lebreton JP, Waite H (eds) (2009) Titan from
Cassini-Huygens. Springer, Dordrecht
Clausen KC et al (2002) The Huygens probe system
Future Directions design. Space Sci Rev 104:155–189
Fulchignoni M et al (2002) The characterisation of Titan’s
atmospheric physical properties by the Huygens atmo-
The lessons learnt from Huygens will be used
spheric structure instrument (HASI). Space Sci Rev
in future to return to Titan and explore in details 104:395–431
its surface in many locations. This was already Fulchignoni M et al (2005) In situ measurements of the
envisaged with an international mission such physical characteristics of Titan’s environment.
Nature 438:785–791
as the Titan Saturn System Mission (TSSM).
Israel G et al (2002) Huygens probe aerosol col-
The TSSM, jointly studied by NASA and ESA, lector pyrolyser experiment. Space Sci Rev 104:
planned to send a lander to explore one of Titan’s 433–468
lake and a Montgolfier to study for several Israel G et al (2005) Complex organic matter in Titan’s
atmospheric aerosols from in situ pyrolysis and analy-
months Titan’s atmosphere and surface.
sis. Nature 438:796–799
Lebreton JP, Matson D (2002) The Huygens probe: sci-
ence payload and mission overview. Space Sci Rev
See Also 104:59–100
Lebreton JP et al (2005) An overview of the descent and
landing of the Huygens probe on Titan. Nature
▶ Aerosols
438:758–764
▶ Atmosphere, Structure Lebreton JP et al (2009) Results from the Huygens probe
▶ Cassini on Titan. Astron Astrophys Rev 17:149–179
▶ Cassini Mission Niemann HB et al (2002) The gas chromatograph mass
spectrometer for the Huygens probe. Space Sci Rev
▶ Cassini-Huygens Space Mission
104:553–591
▶ Chromatography Niemann HB et al (2005) The abundances of constituents
▶ Doppler Shift of Titans’ atmosphere from the GCMS instrument on
▶ GC/MS the Huygens probe. Nature 438:779–784
Owen T (2005) Huygens rediscovers Titan. Nature 438:
▶ Hydrocarbons
756–757
▶ Hydrogen Raulin F, Gazeau MC, Lebreton JP (eds) (2007) A new
▶ Hydrogen Cyanide image of Titan: Titan as seen from Huygens. Planet
▶ Landing Site Space Sci 55(13):1843–2036
Raulin F, Gazeau MC, Lebreton JP (eds) (2008)
▶ Methane
Latest news from Titan. Planet Space Sci 56(5):
▶ Nitrogen 571–777
▶ Pyrolysis GC/MS Tomasko M et al (2002) The descent imager/spectral
▶ Refractory Molecule radiometer (DISR) experiment on the Huygens entry
probe of Titan. Space Sci Rev 104:469–551
Tomasko MG et al (2005) Rain, winds and haze during the
▶ Saturn Huygens probe’s descent to Titan’s surface. Nature
▶ Spectroscopy 438:765–778
▶ Tholins Zarnecki JC et al (2002) Huygens’ surface science pack-
age. Space Sci Rev 104:593–611
▶ Titan
Zarnecki JC et al (2005) A soft solid surface on Titan as
▶ Volatile revealed by the Huygens surface science package.
▶ Water Nature 438:792–795
Hydantoin 1143
▶ Genetics
Huygens Spacecraft ▶ Nucleic Acids
▶ Primer
▶ Huygens Probe ▶ Sequence
▶ Species
Hybridization
Carlos Briones
Hydantoin
Louis d’Hendecourt
Spain
Institut d’Astrophysique Spatiale, Université
Paris-Sud 11, Orsay Cedex, France
Definition
H
Synonyms
In ▶ genetics, hybridization is the process by
which two genetically unrelated
2, 4-Imidazolelidinedione; Glycolyurea
parents – belonging to different strains, varieties,
▶ species, or even genera – are crossed or mated
to originate a hybrid organism. For example, a
Definition
mule is a hybrid animal produced by crossing a
female horse with a male donkey, whereas the
Hydantoin (C3H4N2O2) is an organic compound
grapefruit is a hybrid between a sweet orange and
whose structure is shown in Fig. 1. It is a satu-
a pomelo. In turn, in molecular biology, hybrid-
rated heterocyclic molecule related to imidazole
ization is the process by which two biomolecules
which contains two lactam, or cyclic amide, func-
or fragments of them interact specifically with
tions. From an astrobiological point of view, it
each other. ▶ Nucleic acid hybridization is the
could be important for two main reasons: (1) it is
process of annealing two single-stranded
formed from condensation of ▶ urea and
(ss) DNA or ssRNA molecules of different origin
▶ glycolic acid, or cyanate and ▶ glycine, likely
to form a double-stranded (ds) DNA, a dsRNA, or
common prebiotic molecules which have also
a DNA-RNA duplex. Both nucleic acid strands
been detected in two meteorites, including Mur-
must bear some sequence similarity in order to
chison, and, more importantly, (2) it is a precur-
form stable hybrids by complementary base
sor in the formation of N-carbamoyl amino acid
pairing. The process can be induced in the test
(NCA) or hydantoic acid, which may be an
tube or, alternatively, used for detection and iden-
important intermediate in the formation of
tification of complementary sequences in ex vivo
oligopeptides. More generally, hydantoins are
samples. Different hybridization techniques have
cyclic molecules formed from N-carbamoyl-
been implemented in molecular biology and bio-
amino acids.
technology, including membrane-based blot
methods and microarray technology.
Hydantoin, O
Fig. 1 Hydantoin H
N
See Also
▶ Base Pair O
N
▶ Double Helix H
1144 Hydantoin Formation
variety of small organic compounds including

Hydantoin Formation hydrocarbons as a principal ingredient. The syn-
thesis of hydrocarbon derivatives in the icy
▶ B€
ucherer-Bergs Synthesis mantles of interstellar grains could be initiated
by irradiation by ultraviolet photons from a
remote star.
The hydrocarbon derivatives of meteoritic
origin, once they were delivered to Earth,
Hydrocarbons would mostly be buried in the ocean. A smaller
fraction would eventually be released into the
Koichiro Matsuno periphery regions of hydrothermal environments
Nagaoka University of Technology, Nagaoka, and not necessarily right in the vicinity of black
Japan smokers, which are full of sulfur-containing
chemicals. Insofar as small hydrocarbon deriva-
tives such as formic acid or oxalic acid or their
Keywords precursors are available in the periphery of
hydrothermal environments at temperatures
Carbonaceous meteorites; Fatty acids; Fischer- even less than 200 C, Fischer-Tropsch-type
Tropsch-type reactions; Interstellar grains; Pyru- reactions with the aid of metallic catalysts can
vate; Ultraviolet photon work without being disturbed by the destructive
interference from sulfur (McCollom et al. 1999).
If the reaction solution is maintained at appro-
priately high pressures and temperatures, elon-
Definition
gation of the alkyl chain can occur.
A consequence is the synthesis of saturated
Hydrocarbons are organic compounds consisting
fatty acids such as decanoic acid. Once fatty
entirely of carbon and hydrogen. They may be
acids are synthesized in aqueous environments
synthesized from primary carbon sources such as
where ionic strength is not sufficiently high,
carbon monoxide. The major source of prebiotic
fatty acid vesicles are likely to be formed
hydrocarbons on Earth might have been carbo-
because of their amphiphilic nature. The pres-
naceous meteoritic infalls. Prebiotic significance
ence of the vesicles can provide a means of
of hydrocarbons could be found in their deriva-
encapsulating smaller monomers, enhancing
tives including the carbonyl group.
the likelihood of further synthetic reactions to
proceed in the vesicles.
Hydrocarbon derivatives have also been
Overview synthesized from pyruvate in prebiotic evolution
experiments. Pyruvate itself has been synthe-
A major source of hydrocarbons for prebiotic sized from carbon monoxide using iron sulfide
evolution on Earth may be extraterrestrial mate- working as a reducing agent at 250 C in
rial. Hydrocarbon derivatives constitute several experiments simulating black smokers (Cody
percent of the total mass of carbonaceous mete- et al. 2000).
oritic infalls, in contrast to amino acids that are
present only in trace amounts in carbonaceous
meteorites. The meteoritic infalls are in the See Also
form of interstellar grains composed largely of
silicate minerals coated by thin layers of ice and ▶ Black Smoker, Organic Chemistry
frozen gases of such as carbon dioxide, carbon ▶ Fischer-Tropsch-Type Reaction
monoxide, ammonia, and methanol as well as a ▶ Hydrothermal Reaction
Hydrodynamic Flow 1145
References and Further Reading For a dense atmosphere, gas molecules col-
lide with each other very frequently, and the
Cody GD, Boctor NZ, Filley TR, Hazen RM, Scott JH, distance between successive collisions (mean
Sharma A, Yonder HS Jr (2000) Primordial
free path) is short. Let us imagine traces of
carbonylated iron-sulfur compound and the synthesis
of pyruvate. Science 289:1337–1340 monatomic nitrogen in an atmosphere of mon-
McCollom TM, Ritter G, Simoneit BRT (1999) Lipid syn- atomic hydrogen. Collision with hydrogen,
thesis under hydrothermal conditions by Fischer-Tropsch- which on average tends to expand and leave the
type reactions. Orig Life Evol Biosph 29:153–166
gravity field, imparts momentum to nitrogen
atoms. Although thermal agitation would not
be sufficient to remove this species from an
Hydrodynamic Escape atmosphere of pure nitrogen, the hydrogen vis-
cous drag adds extra velocity to other species
Francis Albarède and may remove them to space. Hydrogen loss
Ecole Normale Supérieure de Lyon, Lyon, France from early planetary atmosphere results in the
loss of other species. This phenomenon, which is
known as hydrodynamic escape, was elucidated H
Definition
by Hunten et al. (1987). There are some sugges-
tions that Mars and Venus may have lost nitro-
The term hydrodynamic escape refers to the
gen and water in this way (e.g., Pepin 1991).
removal of gas from the atmosphere of a planet
possessing a strong gravity field by the viscous
drag of an escaping lighter gas, typically hydro-
See Also
gen. This phenomenon may explain why some
planetary atmospheres are depleted in oxygen,
▶ Earth’s Atmosphere, Origin and Evolution of
nitrogen, and heavier noble gases, such as xenon.

Overview
Hunten DM, Pepin RO, Walker JCG (1987) Mass frac-
When atmospheric temperature is high enough, tionation in hydrodynamic escape. Icarus 69:532–549
the thermal agitation of gas molecules may be Pepin RO (1991) On the origin and early evolution of
strong enough for some gaseous species to leave terrestrial planet atmospheres and meteoritic volatiles.
Icarus 92:2–79
the gravity field of the planet. For a planetary
body with radius Rp, Jean’s escape velocity is
obtained by equating p theffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
kineticffi and potential
energy and is equal to 2GM=Rp , with G being Hydrodynamic Flow
the universal gravitational constant and M the mass
of the planet. For the Earth, escape velocity is Steven B. Charnley
11.2 km s1. The mean thermal (translational)
pffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi Solar System Exploration Division, Code 691,
velocity of atoms in a gas is 8RT=pm , where Astrochemistry Laboratory, NASA Goddard
m is the atomic mass of the gas and R is the gas Space Flight Center, Greenbelt, MD, USA
constant (8.314 103 amu (km/s1)).
For monatomic hydrogen, m = 1 g/mol1 and the
thermal velocity is 2.5 km s1 at ambient temper- Definition
ature (298 K). Although this value is less than
the escape velocity, a tiny fraction of hydrogen The term hydrodynamic flow describes a dynam-
leaves the upper terrestrial atmosphere. In contrast, ical approximation for the bulk motion of matter
the thermal velocity of monatomic nitrogen is in which it is treated as a fluid (gas or liquid),
700 m s1, and nitrogen remains in the atmosphere. rather than as a collection of particles.
1146 Hydrogen
chemistry with many reactions exchanging pro-

Hydrogen tons between soluble molecules.
Molecular hydrogen (H2), which is a gas at
Henderson James (Jim) Cleaves II room temperature and pressure, was first pro-
Earth–Life Science Institute (ELSI), Tokyo duced artificially in the sixteenth century by Par-
Institute of Technology, Meguro–ku, Tokyo, acelsus by mixing metals with strong acids. In
Japan 1766–1781, Cavendish was the first to recognize
Institute for Advanced Study, Princeton, NJ, hydrogen gas as a discrete substance, and that it
USA produces water when burned in air. This property
Blue Marble Space Institute of Science, led to it receiving its name, which is from the
Washington, DC, USA Greek meaning “water-former.” Molecular
Center for Chemical Evolution, Georgia Institute hydrogen is highly inflammable and burns readily
of Technology, Atlanta, GA, USA in air:
2H2 ðgÞ þ O2 ðgÞ ! 2H2 O (l)

Definition
The energy levels of hydrogen can be calculated
Hydrogen (chemical symbol H) is a chemical fairly accurately using the Bohr model of the
element with atomic number 1. It has an atomic atom. However, a more accurate description of
weight of 1.00794 u (or Da) (1.007825 u for the the hydrogen atom comes from a purely quantum
isotope 1H containing no neutrons). It is the mechanical treatment that uses the Schrödinger
lightest and most abundant element, making up equation, which allows the calculation of the
75 % of the Universe’s detectable mass and probability of the electron density around the
90 % of its detectable atoms. proton.
Protonated molecular hydrogen, or H3+, is
found in the interstellar medium, where it is gen-
Overview erated by ionization of molecular hydrogen by
cosmic rays. H3+ is relatively stable in outer
Main sequence stars are mostly composed of space due to the low temperature and matter
ionized hydrogen in plasmas, which plays a density. H3+ is thus one of the most abundant
vital role in powering stars through proton-proton ions in the universe, and plays an important role
reactions and the CNO nuclear fusion cycle. in the chemistry of the interstellar medium.
Hydrogen has three naturally occurring isotopes: Hydrogen forms millions of known molecular
1
H, 2H, and 3H, which all contain 1 proton and compounds with other elements, in particular
0, 1, or 2 neutrons, respectively. While hydrogen carbon. Hydrogen forms molecular compounds
gas is rare in the Earth’s atmosphere (1 ppm by with elements that are more electronegative than
volume) because it escapes from the Earth’s grav- it is, such as the halogens. In these compounds
ity field more easily than heavier gases, it is the hydrogen has a partial positive charge. Hydrogen
third most abundant element on the Earth’s sur- also forms molecular compounds with less elec-
face, mainly as a component of water. tronegative elements, such as the metals and met-
Its most common isotope is 1H (protium), alloids, in which case it takes on a partial negative
which contains a single proton. In ionic com- charge. These compounds are called hydrides.
pounds, hydrogen can take a negative charge Large quantities of hydrogen gas are produced
(forming an anion known as hydride and written by reaction of water with mantle minerals (such
as H), or as a positively charged species H+, also as olivine) notably on the sea floor. It is widely
known as a proton. Hydrogen forms molecular believed that this process, called serpentinization,
compounds with most elements. It plays an may have created a reduced environment favor-
important role in Bronsted-Lowry acid-base able for the origin of life.
Hydrogen Bond 1147
See Also other non-covalent interactions, such as van der

Waals interactions, but weaker than covalent or
▶ Hydrogen Isotopes ionic bonds. The bond strength depends on its
▶ Hydrothermal Environments length, the bond angle, the local dielectric con-
▶ Serpentinization stant, the electronegativity of the non-hydrogen
atoms, temperature, and pressure.
Hydrogen bonds can occur between molecules
(intermolecular hydrogen bond) or within differ-
ent parts of a single molecule (intramolecular
hydrogen bond).
Hydrogen Bond The most ubiquitous example of an
intermolecular hydrogen bond is the one found
Gilles Bruylants
between water molecules. The water molecule is
Engineering of Molecular NanoSystems,
composed of two hydrogen atoms which can
Universté Libre de Bruxelles, Brussels, Belgium
form a hydrogen bond (as donors) and one oxy-
gen atom which has two lone pairs of electrons, H
each of which can be involved in a hydrogen
Keywords
bond (as acceptors). In liquid water, every water
molecule can consequently be hydrogen bonded
Molecular interactions; Noncovalent interactions
with up to four other molecules.
Hydrogen bonding also plays an important
role in determining the three-dimensional struc-
Synonyms
ture adopted by larger molecules including mac-
romolecules such as ▶ proteins and ▶ nucleic
H bond
acids. The double helical structure of DNA is
due largely to hydrogen bonding between the
▶ base pairs, which link one complementary
Overview strand to the other. In proteins, hydrogen bonds
between the backbone oxygen atoms and amide
A hydrogen bond (H bond) is the attractive hydrogen atoms contribute to the formation of
non-covalent interaction between a hydrogen secondary structure elements, such as alpha heli-
atom covalently bonded to an electronegative ces and beta sheets. In addition, many proteins
atom (hydrogen bond donor) and another electro- have hydrogen bonds between side chains. Taken
negative atom (hydrogen bond acceptor), such as together, these interactions contribute to folding
nitrogen, oxygen, or fluorine. proteins into specific shapes that determine their
The hydrogen bond has some features of cova- biological function. However, the importance of
lent bonding: it is directional, produces this contribution to stability of the folded state
interatomic distances shorter than the sum of the still remains controversial, as both donors and
van der Waals radii, and usually involves a lim- acceptors of hydrogen bonds can form hydrogen
ited number of interacting partners, which can be bonds with water when unfolded in aqueous
interpreted as a type of valence. These covalent solution.
features are more substantial when the hydrogen
atom is bonded to a more electronegative atom.
The hydrogen bond is relatively strong (usually See Also
between 5 and 30 kJ/mol but extremely strong
values such as 155 kJ/mol in the HF2 molecule ▶ Base Pair
are observed, in which case the term Low Barrier ▶ Covalent Bonds
Hydrogen Bond is frequently used) compared to ▶ Nucleic Acids
1148 Hydrogen Chloride
▶ Protein See Also

▶ Van der Waals Forces
▶ Water, Solvent of Life ▶ Chlorine Hydrides in the Interstellar Medium
▶ Watson-Crick Pairing ▶ Interstellar Dust
▶ Weak Bonds ▶ Molecular Cloud
Atkins P, de Paula J (2009) Atkins physical chemistry,
9th edn. Oxford University Press, Oxford, New York Blake GA, Keene J, Phillips TG (1985) Chlorine in dense
interstellar clouds – the abundance of HCl in OMC-1.
Astrophys J 295:501
Cernicharo J, Goicoechea J, Daniel F et al (2010a) The
35
Cl/37Cl isotopic ratio in dense molecular clouds:
HIFI observations of hydrogen chloride towards
Hydrogen Chloride W3A. Astro Astrophys 518:L115
Cernicharo J, Decin L, Barlow M et al (2010b) Detection
William M. Irvine of anhydrous hydrochloric acid, HCl, in IRC+10216
with the Herschel SPIRE and PACS spectrometers.
University of Massachusetts, Amherst, MA, USA Astron Astrophys 518:L136
Cernicharo WR, Decin J et al (2010c) A high resolution
line survey of IRC+10216 with Herschel First results:
Synonyms detection of warm silicon dicarbide (SiC2). Astron
Astrophys 521:L8
De Luca M, Gupta H, Neufeld D, Gerin M, Teyssier D,
HCl Drouin BJ, Pearson JC, Lis DC, Monje R, Phillips TG,
Goicoechea JR, Godard B, Falgarone E, Coutens A,
Bell TA (2012) Herschel-HIFI Discovery of HCl+ in
the interstellar medium. Astrophys J 751:L37–L42
Definition Neufeld DA, Roueff E, Snell RL et al (2012) Herschel
observations of interstellar chloronium. Astrophys
Hydrogen chloride is a colorless gas under stan- J 748:37–47
dard conditions, consisting of diatomic mole- Schilke P, Phillips TG, Wang N (1995) Hydrogen chloride
in OMC-1. Astrophys J 441:334–342
cules containing hydrogen and chlorine. It is
detected in the gas phase in interstellar molecular
clouds (Schilke et al. 1995), but the majority of
the chlorine in these regions appears to be seques- Hydrogen Cyanamide
tered in the ▶ interstellar dust grains (Cernicharo
et al. 2010a). The two isotopic species, H35Cl and ▶ Cyanamide
H37Cl, have been detected with the Herschel sat-
ellite toward evolved stars (Cernicharo
et al. 2010b, c). More recently, the cations HCl+
Hydrogen Cyanide
and H2Cl+ have also been detected in molecular
clouds (e.g., De Luca et al. 2012; Neufeld
et al. 2012).
Molecular Astrophysics, Iorrejón de Ardoz,
Madrid, Spain
History
Interstellar HCl was first detected in 1985 from Synonyms

the Kuiper Airborne Observatory by G. Blake and
colleagues. Formonitrile
Hydrogen Isocyanide 1149
Definition Saykally RJ, Szanto PG, Anderson TG, Woods RC

(1976) The microwave spectrum of hydrogen
isocyanide. Astrophys J 204:L143
Due to its dual reactivity, as both a nucleophile Snyder LE, Buhl D (1971a) Observations of radio emis-
and an electrophile, hydrogen cyanide, HCN, sion from interstellar hydrogen cyanide. Astrophys
plays an important role in many synthetic pro- J 163:L47
cesses potentially related to the origin of life. At Snyder LE, Buhl D (1971b) Bull Am Astr Soc 3:388
Wilson WJ, Schwartz PR, Epstein EE (1973) Carbon mon-
room temperature, it is a colorless and extremely oxide and hydrogen cyanide millimeter wave emission
poisonous gas, with a bitter odor reminiscent of from stars. Astrophys J 183:871
almonds that some people are unable to perceive
for genetic reasons. It has been found in molecu-
lar clouds (Snyder and Buhl 1971), evolved stars
(Wilson et al. 1973), and comets (Huebner
et al. 1974; Despois et al. 1986). It is one of the Hydrogen Cyanide Polymer
most abundant triatomic species in space. Its less
stable isomer ▶ hydrogen isocyanide, HNC, has ▶ HCN Polymer
also been found in space (Snyder and Buhl 1971; H
Saykally et al. 1976). In molecular clouds, both
isomers form from the dissociative electronic
recombination of HCNH+. In cold molecular
clouds, the abundance ratio HCN/HNC is of Hydrogen Ion Concentration Index
order unity (Sarrasin et al. 2010). HCN plays an
important role in chemical reactions thought to ▶ pH
have been important on the primitive Earth, for
example, in Miller-Urey-type experiments (via
its involvement in the Strecker amino acid syn-
thesis) and in its self-condensation reactions that
produce both amino acids and nucleic acid bases Hydrogen Isocyanide
(Oró 1961).
William M. Irvine
See Also
Synonyms
▶ HCN Polymer
▶ Hydrogen Isocyanide
HNC
▶ Miller, Stanley
Definition
Hydrogen isocyanide (HNC) is a linear triatomic
Despois D, Crovisier J, Bockelee-Morvan D, Gerard E, molecule that is an isomer of HCN, ▶ hydrogen
Schraml J (1986) Observations of hydrogen cyanide in
cyanide, the latter being a key intermediary in
comet Halley. Astron Astrophys 160:L11
Huebner WF, Buhl D, Snyder LE (1974) HCN radio several possible pathways for the production of
emission from Comet Kohoutek/1973f/. Icarus 23:580 important biological molecules. HNC itself is
Oró J (1961) Mechanism of synthesis of adenine from one of the more abundant triatomic molecular
hydrogen cyanide under possible primitive earth con-
species in interstellar clouds, where the abun-
ditions. Nature 191:1193
Sarrasin E, Abdallah DB, Wernli M, Faure A, dance ratio HNC/HCN appears to be sensitive to
Cernicharo J, Lique F (2010) MNRAS 404:518 temperature. Because of its relatively large
1150 Hydrogen Isotopes
electric dipole moment (and hence rapid transi-

tions to lower energy states) and since higher Hydrogen Isotopes
energy levels are populated by collisions with
molecular hydrogen (the most abundant cloud Daniele L. Pinti
species), the intensity of HNC rotational transi- GEOTOP Research Center for Geochemistry and
tions can be used to estimate density in molecu- Geodynamics, Université du Québec à Montréal,
lar clouds. Hydrogen isocyanide is also observed Montréal, QC, Canada
in cometary comae.
History Definition
Hydrogen isocyanide was initially detected in Hydrogen (symbol, H; atomic mass, 1.00794) has
the interstellar medium in 1971 by L. Snyder three isotopes. Two are stable in natural environ-
and D. Buhl. Because HNC is considerably ments and are denoted as 11H and 21H (deuterium or
higher in energy than its isomer HCN, at thermal D). The third isotope 31H (tritium or T) is unstable
equilibrium its abundance in the interstellar and decays to 3He with a ▶ half-life of
medium would be essentially zero. Instead, in 12.32 years. Being hydrogen one of the
cold clouds, its abundance can equal or even ▶ water-forming elements, its isotopic ratio D/H
exceed that of HCN. This evidence for the has been used for identifying the origin of water
kinetic control of isomerization pathways was within the Solar System. Particularly, the chon-
one of the strongest early observational supports dritic D/H ratio measured in meteorites and Ant-
for ion-molecule models of interstellar chemis- arctic micrometeorites (143–170 106) is
try (Herbst 1987). Hydrogen isocyanide was closer to the value of the bulk Earth
first observed in a comet in Comet Hyakutake (149–153 106) and the present-day ocean
in 1996, but there is evidence that it may be (155.7 106) than the comets
primarily produced in the coma rather than (298–324 106). This is the isotopic evidence
being a constituent of the icy nucleus (Irvine often used for claiming that water on Earth has
et al. 1998). been delivered by chondritic planetesimals rather
than comets.
See Also
See Also
▶ Comet
▶ Delta, Isotopic
▶ Molecular Cloud
▶ Deuterium/Hydrogen Ratio
▶ Micrometeorites
▶ Oxygen Isotopes
References and Further Reading ▶ Water
Herbst E (1987) Gas phase chemical processes in molec-
ular clouds. In: Hollenbach DJ, Thronson HA (eds)
Interstellar processes. D. Reidel, Dordrecht,
pp 611–630
Irvine WM, Bergin EA, Dickens JE, Jewitt D, Lovell AJ,
Matthews HE, Schloerb FP, Senay M (1998) Chemical
Hydrogen Oxide
processing in the coma as the source of cometary HN-
C. Nature 393:547–550 ▶ Water, Solvent of Life
Hydrogenosomes 1151
Definition
Hydrogen Phosphide
Amorphous carbon is composed of randomly
▶ Phosphine arranged clusters of hexagonal rings (the carbon
portion of benzene). When hydrogen atoms are
attached at surface sites, the substance is referred
Hydrogen Sulfide to as hydrogenated amorphous carbon or
HAC. At various times, hydrogenated amorphous
Nicholas Arndt carbon has been proposed as the carrier of the
ISTerre, Université Grenoble Alpes, France infrared emission features commonly associated
with ▶ polycyclic aromatic hydrocarbons.
Definition
See Also
Hydrogen sulfide is a colorless and poisonous
gas with a strong odor of rotten eggs and the ▶ Polycyclic Aromatic Hydrocarbon H
formula H2S. It occurs in volcanic gases and ▶ Unidentified Infrared Emission Bands
natural gas and is produced by the breakdown
of organic matter, commonly through the action
of bacteria, and by liberation from sulfide min-
erals. H2S figures prominently in many origins of
Hydrogenated Fullerene
life models, especially those positing an origin
near ▶ black smoker-type hydrothermal vents.
▶ Fullerane
It is found both in interstellar molecular clouds
and in ▶ comets. The sulfanyl radical, SH,
and the corresponding cation sulfaniumylidene
(sulfanylium), SH+, have also recently been
Hydrogenosomes
detected in the interstellar medium
(▶ Molecules in Space).
Ricardo Amils
See Also Universidad Autónoma de Madrid, Madrid, Spain
▶ Black Smoker
▶ Comet
Definition
▶ Sulfur Hydrides in the Interstellar Medium
Hydrogenosomes are organelles present in
certain anaerobic eukaryotic microorganisms
in which ▶ fermentation reaction takes place to
generate ▶ ATP. Some anaerobic eukaryotes
Hydrogenated Amorphous Carbon lack mitochondria and contain instead
hydrogenosomes. The major biochemical reac-
William M. Irvine
tions in the hydrogenosomes are those associ-
ated to the oxidation of pyruvate producing H2,
CO2, and acetate as final products. In some
Synonyms anaerobic ▶ eukaryotes, intracellular symbiotic
H2 consumers, such as ▶ methanogens, can be
HAC found.
1152 Hydrolysis
See Also products are low (on the order of 103–106 molar),
the hydrolysis reaction, such as the hydrolysis of a
▶ ATP peptide to amino acids, is essentially thermody-
▶ Endosymbiosis namically irreversible. For example:
▶ Energy Conservation
▶ Eukaryote Glycylglycine þ H2 O ! 2 Glycine
▶ Fermentation
▶ Methanogens
Here the DG for the reaction is 3.5 kcal/mole,
and the spontaneous reassembly of amino acids to
▶ peptides is negligible.
Hydrolysis
See Also
Earth–Life Science Institute (ELSI), Tokyo ▶ Amide
Institute of Technology, Meguro–ku, Tokyo, ▶ Amine
Japan ▶ Carbonyl
Institute for Advanced Study, Princeton, NJ, ▶ Carboxylic Acid
USA ▶ Peptide
Washington, DC, USA
Hydrophobic Effect
Lawrence Pratt
Definition Tulane University, New Orleans, LA, USA
In chemistry, hydrolysis refers to a chemical

reaction in which molecules are split by the addi- Keywords
tion of a molecule of water. One fragment of the
parent molecule gains a hydrogen ion (H+), while Hydrophobicity; Hydrophobic interactions;
the other group receives a hydroxyl group (OH). Hydrophobic bonds
Hydrolytic reactions catalyzed by both acid
and base are common, for example, the hydroly-
sis of ▶ amides or esters. Their hydrolysis occurs Definition
when a nucleophile (water or a hydroxyl anion)
attacks the ▶ carbonyl group of the ester or Hydrophobic effects are phenomena of aqueous
amide. In water, hydroxyl ions are better nucleo- solutions involving hydrophobic species or
philes than water itself. In acid, the carbonyl molecular groups, which are recognized by con-
group becomes protonated, and this leads to eas- trast to hydrophilic species or groups. Hydro-
ier nucleophilic attack. The products of the philic species typically have simply recognized
hydrolysis of amides and esters are a carboxylic chemical interactions with water molecules, e.g.,
acid and an amine, and a carboxylic acid and an ammonia (NH3) or perhaps have electrostatic
alcohol, respectively. interactions with water, e.g., ions such as Na+.
Under physiological conditions, in which the Hydrophilic species typically have substantial
concentrations of a hydrolysable reactant and its solubility in liquid water. In contrast,
Hydrophobic Effect 1153
Hydrophobic Effect, Fig. 1 Ostwald coefficient L for of O2 in the water is 5 % of the concentration of O2 in
some simple gases in water (circles) and cyclohexane coexisting vapor. All results are for low (Earth) ambient
(stars). The Ostwald coefficient is the fractional amount pressures. Notice that this plot utilizes a logarithmic ver- H
of the gas in the solution relative to the amount in a tical axis. These gases are less soluble in water than in
coexisting dilute vapor phase. Thus L = 0.05 for O2 at cyclohexane, and the T dependences display opposite
the lower temperatures here means that the concentration trends (Redrawn after Ashbaugh and Pratt (2006))
hydrophobic species, e.g., methane (CH4) hydrophobic and hydrophilic effects can make
or hydrocarbons broadly, have less favorable the solution thermodynamic characterization of
interactions with water molecules, and predomi- these solutions tricky. Since hydrophilic effects
nantly interactions of van der Waals type. Hydro- are typically more simply recognized and
phobic species typically have low solubilities in modeled, hydrophobic effects are associated
water. Hydrophobic groups can be combined with the additional complications exhibited by
with hydrophilic groups in larger molecules, amphiphilic species. Those complications are
and this vastly extends the functionality of typically unfavorable entropy change for disso-
molecules like that. Molecules that combine lution, and substantial temperature dependences
hydrophobic and hydrophilic groups are for those entropy changes. Figure 1 gives exam-
called amphiphilic. Sodium dodecylsulfate ples of distinctions between solubilities of sim-
(CH3(CH2)11OSO3Na), a simple surfactant, ple gases in water and cyclohexane, and
is an example. The characteristic functionality distinctions in T-dependences. Distinctive
of amphiphilic molecules is to coat aqueous solu- T-dependences are often required for identifica-
tion interfaces, satisfying the tendency tion of hydrophobic effects for specific study.
of hydrophilic groups for contact with water
while also satisfying the tendency of hydropho-
bic groups to avoid contact with water. History
In fact, the appellation “surfactant” indicates
that behavior. Tanford’s (1997) expansive discussion (Further
The extended functionality of amphiphilic spe- Reading) gives an entertaining history of the gen-
cies can lead to self-assembled possibilities for esis of the concepts and names associated with
organization of larger solution structures, includ- hydrophobic effects in the context of understand-
ing micelles, bilayer membranes, and specifically ing soluble proteins. That historical discussion
structured macromolecules such as folded water- culminates with the watershed review article of
soluble protein molecules. The balance of Kauzmann (1959) (Further Reading), which
1154 Hydrophobic Effect
marks an opening of the modern era in study of the solution in stabilizing functional biomolecular
hydrophobic effects. Since the mid-1970s, anal- structures is an aspect of hydrophobic effects.
ysis of the underlying molecular description Molecular theories and explanations of hydro-
of hydrophobic effects has extensively exploited phobic effects range from appeal to and parame-
the development of molecular-scale simulation terization of simple intuition to aggressive
with rapidly advancing computational tools. statistical mechanical theory of solutions. Because
Ashbaugh and Pratt (2006) presented a modern these are problems of acknowledged importance
perspective on the interplay between computer and sometimes challenging to molecular-scale
simulation work, molecular theory, and classic intuition, agreement on molecular theories and
physical chemistry. A broader range of applica- mechanisms has been imperfect. Ashbaugh and
tions is considered in the review of Pratt and Pratt (2006) give a glimpse of the range molecular
Pohorille (2002). theories advanced for these phenomena.
An important aspect to hydrophobic effects is
that the solvent water itself behaves unlike com-
Overview mon organic solvents. Liquid water is typically a
chemically active medium. It is stable over com-
A striking feature of observed biophysical struc- paratively wide temperature range, and exhibits
tures is that spatial organization is achieved by a large heat capacity that helps to moderate tem-
competing interactions, hydrophilic and hydro- perature excursions for the terrestrial biosphere.
phobic interactions. The most primitive concep- The Pohorille and Pratt (2012) discussion con-
tualization of hydrophobic effects is just that sidered thermodynamic characteristics of sol-
water and oil can demix. Those phase equilibria, vents that might be compared to water. That
together with molecular-scale hydrophobic- survey identified non-water liquids glycerol,
hydrophilic amphiphilicity, can then result in diethylene glycol, ethylene glycol, formamide,
spatially segregated mesoscopic structures 2-aminoethanol, 1–4 butanediol, and 1–5
(Further Reading: Tanford 1978). The driving pentanediol, and mixtures of such solvents, as
force for oil-water phase separation is customar- cases worthy of further study for comparison
ily viewed as a sticky interaction operating with liquid water in supporting a solvophobic/
between hydrophobic species, but not as specifi- solvophilic dichotomy and perhaps leading to
cally as suggested by the old-fashioned term interesting self-assembly of functional larger-
“hydrophobic bond”. scale solution structures.
Phase separation is not the only interesting
aspect of hydrophobic effects. Standard solution
thermodynamic studies establish that the See Also
suggested hydrophobic stickiness becomes stron-
ger as the temperature increases through a physio- ▶ Lipid Bilayer
logical range. This is experimentally clear in the ▶ Protein
phenomenon of cold-denaturation of proteins ▶ Water
wherein cool, unfolded soluble proteins fold upon
heating. That protein globules become more References and Further Reading
highly ordered with increasing T may seem
counter-intuitive. But if the contacting aqueous Ashbaugh HS, Pratt LR (2006) Colloquium: scaled parti-
solution becomes sufficiently disordered with cle theory and the length scales of hydrophobicity. Rev
T-increase, expectations from ordinary thermody- Mod Phys 78:159–178
Kauzmann W (1959) Some factors in the interpretation of
namics are restored. This suggests that the protein denaturation. Adv Protein Chem 14:1–63
refolding of a cold-denatured protein is driven by Pohorille A, Pratt LR (2012) Is water the universal solvent
the participation of the solution. Participation of for life? Orig Life Evol Biosph 42:405–409
Hydrosphere 1155
Pratt LR, Pohorille A (2002) Hydrophobic effects and sometimes used to evaluate the hydrophobicity
modeling of biophysical aqueous solution interfaces. of amino acids and other compounds.
Chem Rev 102:2671–2692
Pratt LR, Pohorille A, Asthagiri D (2007) What is special
about water as a matrix of life? arXiV;physics/
0701282 See Also
Tanford C (1978) The hydrophobic effect and the organi-
zation of living matter. Science 200:1012–1018
Tanford C (1997) How protein chemists learned about the ▶ Amino Acid
hydrophobic factor. Protein Sci 6:1358–1366 ▶ Amphiphilicity
▶ Hydrogen Bond
▶ Self-Assembly
Hydrophobicity
Kensei Kobayashi Hydrosphere

Hodogayaku, Yokohama, Japan H
Nicholas Arndt
Definition
Definition
Hydrophobicity refers to the property of a mol-
ecule to resist dissolution in water. Organic com- The hydrosphere is the combined mass of water
pounds with ionizable or polarizable groups tend in the atmosphere, at the surface, and in the
to be hydrophilic, as opposed to hydrophobic, as interior of the Earth and of other planetary bod-
those groups may be favorably solvated by polar ies. About 97 % of the total volume of the
water molecules via hydrogen bonds and other Earth’s hydrosphere (ca. 1.34 109 km3) is in
mechanisms. On the other hand, organic com- the oceans; the remaining 3 % is in the conti-
pounds without polarizable groups are generally nental ice caps (1.7 %), groundwater (1.3 %),
hydrophobic and thus show low solubility in lakes and rivers (0.013 %), and the atmosphere
water. Alkanes are typical hydrophobic mole- (0.0009 %). Up to five times the surface vol-
cules. When hydrophobic molecules are placed ume of the oceans may be present in the mantle,
in water, water tends to exclude them. This is fixed in low concentrations in the structure of
called the hydrophobic effect. Molecules with nominally anhydrous minerals. The near-sun
both hydrophilic groups and hydrophobic orbit of the Earth, well inbound of the crystalli-
groups are called amphiphilic molecules. zation temperature of water, suggests that the
Hydrophobicity and amphiphilicity are impor- Earth initially formed as a rather dry body and
tant in the self-assembly of organic molecules that most of the Earth’s water was likely subse-
which may have been important in the origin of quently supplied to the Earth by meteorites and
biological membranes. cometary material soon after the Moon-forming
Amino acids show a variety of hydrophobic- impact (▶ Late Veneer). Mars appears to have
ities due to their variable side chains. Phenylala- had a small liquid hydrosphere early in its his-
nine and leucine have large hydrocarbon groups tory and retains water in ice caps and in the
and show high hydrophobicity, while aspartic subsurface. A thick hydrosphere covers the
acid and lysine show low hydrophobicity (high Jovian moons Europa (ca. 3.8 109 km3) and
hydrophilicity) due to ionizable groups in their perhaps Ganymede and the Saturnian moon
side chains. Various hydrophobic scales are Enceladus.
1156 Hydrostatic Balance
See Also
Hydrothermal Alteration
▶ Water, Delivery to Earth Michel Jébrak
Département des Sciences de la Terre et de
l’Atmosphère, Université du Québec à Montréal,
Montreal, QC, Canada
Hydrostatic Balance
Keywords
▶ Hydrostatic Equilibrium
Hydrothermal system; Mars
Hydrostatic Equilibrium
Synonyms
Daniel Rouan
Metasomatism
Meudon, France
Definition
Synonyms
Hydrothermal alteration is defined as any change
in the mineralogic composition of a rock due to
Hydrostatic balance
the action (by either physical or chemical means)
of hydrothermal fluids in an open system.
Definition
History
Hydrostatic equilibrium designates a stable state of
a fluid where there is a condition of balance
Hydrothermal alteration has been recognized on
between gravitational force and pressure gradient.
Earth since the onset of geology. It is a very
In a planetary atmosphere, under simplifying
common process, and it is often associated with
assumptions (perfect gas, isothermal), the hydro-
mineral deposits. Hydrothermal alteration was
static equilibrium condition leads to a law of pres-
suspected on Mars from observations carried
sure (p) variation with altitude (z) that is
out by early ▶ Viking landers (Newson and
exponential: p = p0 exp(z/z0), where z0 is the
Hagerty 1997), and it was discovered on the
scale height (8 km on Earth). In a star, the radiation
ground by the Mars Science Laboratory rover
pressure creates an additional outward force that
“Curiosity.”
participates in the hydrostatic equilibrium equation.
Note that hydrostatic equilibrium implies that the
star’s or planet’s shape is a sphere in the simplest
Overview
case and an oblate spheroid for a rotating body.
Hydrothermal alteration marks the disequilib-
rium between fluids, usually aqueous, and rocks,
See Also usually silicate-rich. Hydrothermal alteration has
been observed on Mars and is presumed to exist
▶ Atmosphere Escape on other small ocean planets (Pirajno 2009).
▶ Gravitation Smaller planets show rock porosity to a greater
Hydrothermal Environments 1157
depth than the Earth, allowing deeper hydrother- References and Further Reading
mal fluid circulation and exchange to exist.
On ▶ Mars, alteration mineralogy can be Ehlmann BL, Mustard JF, Murchie SL, Bibring JP,
Meunier A, Fraeman AA, Langevin Y (2011) Subsur-
determined from the composition of impact cra-
face water and clay mineral formation during the early
ters, which provide natural probes into the sub- history of Mars. Nature 479:53–60
surface. Alteration minerals occur as large Michalski JR, Cuadros J, Niles PB, Rogers AD, Wrigth SP
unsaturated, partially altered zone, without con- (2013) Groundwater activity on Mars and implications
for a deep biosphere. Nat Geosci 6:133–138
tact with the atmosphere. Mud volcanoes reflect
Newton HE, Hagerty JJ (1997) Chemical components of
the importance of such clay formation at depth. the Martian soil: melt degassing, hydrothermal alter-
Two styles of alteration have been recorded: ation, and chondritic debris. J Geophys Res
low-pH alteration, marked by sulfate deposits, 102:19345–19355
Pirajno F (2009) Hydrothermal processes and mineral
and neutral to high-pH alteration, marked by
system. Springer, Berlin, 1250 p
crystallization of chlorite, nontronite, saponite, Vance S, Harnnmeijer J, Kimura J, Hussmann H,
serpentine, and carbonates. Such alteration Demartin B, Brown M (2007) Hydrothermal systems
could be produced by hydrothermal systems in small ocean planets. Astrobiology 7:987–1005
related to impacts, high-temperature geothermal H
systems related to plume volcanism (Tharsis),
and zones of high geothermal gradient
(Ehlmann et al. 2011). Impact activity induced Hydrothermal Environments
localized hydrothermal system near the borders
of the craters, and such activity may last several Nicholas Arndt
million years after the impact. The subsurface ISTerre, Université Grenoble Alpes, France
alteration peaked in the early stage of Mars’
evolution, in the Noachian (>4.1 Gyr ago) and
into the Early Hesperian (3.7–4.1 Gyr ago). Ser- Keywords
pentine is uncommon but has been found in
mélange terrains, impact craters, and within a Black/white smoker; Hydrothermal fluid; Oce-
regional olivine unit near the Isidis basin. anic crust; Origin of life
In other small ocean planets (▶ Titan, ▶ Gan-
ymede, ▶ Callisto, ▶ Europa, ▶ Enceladus),
hydrothermal alteration is assumed due to the Definition
probable onset of hydrothermal systems and the
formation of liquid water at the contact within the A hydrothermal environment is a setting domi-
cryosphere (Vance et al. 2007). nated by the circulation of hot, mainly aqueous
Zones of hydrothermal alteration may impact fluids. In a mid-ocean ridge setting, seawater
the topography by facilitating collapse of hydro- penetrates the crust, becomes heated, interacts
thermal zones. Increase of porosity, higher tem- with the crust so that its composition changes,
perature, and presence of water in hydrated then exits the crust at a hydrothermal vent on the
minerals may have facilitated the establishment ocean floor. The environment surrounding the
of subsurface biota. Carbon monoxide-bearing vent sustains rich, anoxic chemotrophic ecology.
fluids infiltrated from the surface could have This environment is a possible setting for the
reacted with the hydrogen from serpentinization emergence of life on Earth.
to form methane, which could have fuelled
▶ methanotrophs and been mediated by
▶ methanogens (Michalski et al. 2013). Alter- Overview
ation of basaltic rocks releases Fe and Mg cations
which can fuel iron respiration and facilitate the The simple definition of a hydrothermal fluid is a
formation of complex organic molecules. hot water-rich fluid in the Earth’s crust. The
1158 Hydrothermal Environments
origin of the fluid is diverse, ranging from reactions between seawater and ultramafic rocks
seawater that penetrates down into the oceanic (▶ serpentinization) produce methane- and
crust or fills pore space in sedimentary basins hydrogen-rich fluids that are highly alkaline,
to magmatic fluid released from crystallizing with lower temperatures ranging from about
igneous intrusions. These fluids are heated 40 C to 90 C. These fluids are rich in dissolved
as they descend into the crust or come into carbonate and sulfate; when they exit from the
contact with magmas. Hot fluids are capable ocean floor, they form “white smokers.”
of transporting large volumes of material The areas around hydrothermal vents often
derived from the parent magma or leached host complex communities of archaea, bacteria,
from surrounding rocks, and many of the world’s and larger, more diverse organisms, including
greatest ore deposits form as metals such as giant tubeworms, clams, limpets, and shrimp.
Cu, Zn, Au, or U precipitate from hydrothermal These deep-sea organisms have no access to
fluids. sunlight and depend on the hydrothermal fluids
In the astrobiological context, the most for nutrients and energy. The archaea and bacte-
important hydrothermal environment is close to ria use sulfur compounds, particularly hydrogen
the surface of the oceanic crust, at or near a sulfide, to produce organic matter via chemosyn-
mid-ocean ridge. As seawater penetrates the thesis. The bacteria surround the hydrothermal
oceanic crust, it becomes heated and interacts vent area with a thick mat that attracts other
with the crust so that its chemical composition organisms such as amphipods and copepods
changes (Humphris et al. 1995; Alt and Teagle that graze on the bacteria. Larger animals com-
2000; Fr€ uh-Green et al. 2007). It then may exit plete the food chain. The main families found
the crust at a hydrothermal vent on the ocean around seafloor vents are annelids, pogonopho-
floor. Components leached from basaltic rocks rans, gastropods, bivalves, and crustaceans,
of the crust – sulfide or sulfate, silica, oxides, and together with the famous tubeworms that epito-
carbonates – precipitate as the fluid is cooled and mize the vent community.
becomes diluted as it mixes with seawater. Corliss et al. (1981) and Wächtershäuser
These components are initially dispersed in the (1990), a German chemist turned patent lawyer,
plume that ascends from the vent, or they floc- suggested that life might have originated at
culate around the vent to form chimneys whose hydrothermal vents early in the Archean. It is
height might reach 60 m. Particles raining out well established that organic compounds are
from the plume produce a layer of sediment, to produced by inorganic, Fischer-Tropsch-type
which the relicts of chimneys (which topple reactions deep in the oceanic crust when hot
when they become too tall and unstable) are aqueous fluids react with minerals like olivine
added. The product is a lens of “exhalative” or pyroxene. These compounds were transported
sulfide, which, if accreted to the continent, can in hydrothermal fluids to the seafloor where they
be mined for metals such as Cu, Zn, and Pb interacted with seawater. The presence of clay
(Tivey 2007). minerals provided templates that foster the
Fluids that circulate at a mid-ocean ridge formation of peptides and protocells. Martin
become strongly heated and acidic as they inter- and Russell (2007) proposed that as hydrother-
act with basaltic crustal rock. Such fluids dis- mal fluid emerged at the ocean floor, it reacted
solve large quantities of Fe-Cu-Zn sulfides that inside small metal sulfide cavities within hydro-
precipitate as small particles within the fluid as thermal vent chimneys. Large chemical and
it exits the vent, imparting a black color to the thermal gradients between the interior and exte-
hydrothermal plume and giving rise to the term rior of the chimney provided the microenviron-
“▶ black smoker.” Such fluids interact with very ment for life-forming chemical reactions to
hot rock adjacent to magma chambers beneath take place.
the ocean ridge; their temperatures may reach Active hydrothermal vents may exist on
400 C. In settings off the mid-ocean ridge axis, Jupiter’s moon Europa, and alteration minerals
Hydrothermal Environments 1159
that might have resulted from the circulation of other oceanographic programs. The biological
hydrothermal fluids have been found on Mars. community is involved through their research
on the ecology of hydrothermal vents. The
search for a better understanding of the reactions
Basic Methodology that produced prebiotic compounds continues.
Companies and government organizations are
Active hydrothermal vent systems are observed investigating how active submarine hydrother-
from submersibles – small submarines that can mal systems might be mined as a source of
descend to the great depths of the ocean floor. metals such as Cu, Zn, and Au.
Scientists record the activity of the vents, regis-
ter the flux and the physical characteristics of the
fluids, and take samples of the fluids, of the
chimneys, and of surrounding sedimentary See Also
deposits. These are analyzed in the
▶ Black Smoker
laboratory – the fluids for their composition
and other characteristics (acidity, oxidation ▶ Chemotroph H
state), the rock samples for their mineralogy,
▶ Origin of Life
and geochemical composition. Several vents
have been sampled by scientific drilling as part ▶ Serpentine
▶ Serpentinization
of the Integrated Oceanic Drilling Program
▶ White Smoker
(IODP). Fossil hydrothermal vents are studied
in sections that have been obducted onto, or
accreted to, the continental crust. Notable exam-
ples are on Cyprus and in mining areas such as References and Further Reading
the Abitibi belt in Canada and the Urals.
Alt JC, Teagle DAH (2000) Hydrothermal alteration
and fluid fluxes in ophiolites and oceanic crust.
In: Dilek Y, Moores E, Elthon D, Nicolas A (eds)
Key Research Findings Ophiolites and oceanic crust: new insights from field
studies and the ocean drilling program, vol 349. Geo-
logical Society of Amercia Special Paper, Boulder,
Hydrothermal circulation plays an important role p 273–282
in cooling the oceanic crust; hydration of basalts Corliss JB, Baross JA, Hoffman SE (1981) A hypthesis
strongly affects its composition; release of fluids concerning the relationship between submarine hot
in subduction zones controls island arc spring, and the origin of life on Earth. Oceanol Acta
1:59–69
magmatism and contributes directly to the growth Fr€
uh-Green GL, Delacour A, Boschi C, Bernasconi SM,
of continental crust. Hydrothermal fluids form Butterfield D, Kelley DS (2007) Building lost city:
accumulations of Cu, Zn, and Pb sulfides, which serpentinization, mass transfer and life in a
are mined when the deposits are found on the peridotite-hosted hydrothermal system. Geochim
Cosmochim Acta 71:A298–A298
continents. Unusual bacterial and animal com- Humphris SE, Zierenberg RA, Mullineaux LS, Thomson
munities surround active hydrothermal events. RE (1995) Seafloor hydrothermal systems: physical,
Hydrothermal vents in the early Archean may chemical, biological, and geological interactions,
have been the site of the emergence and early AGU monograph series, no 91. American Geophysical
Union, Washington, DC
evolution of life. Martin W, Russell MJ (2007) On the origin of biochemis-
try at an alkaline hydrothermal vent. Phil Trans R Soc
Lond B 362:1887–1925
Future Directions Tivey MK (2007) Generation of seafloor hydrothermal
vent fluids and associated mineral deposits. Oceanog-
raphy 20(1):50–65
Investigation of hydrothermal vents is continu- Wächtershäuser G (1990) Evolution of the first metabolic
ing as part of the Ocean Drilling Program and cycles. Proc Natl Acad Sci 87:200–204
1160 Hydrothermal Reaction
reactions examined under the present methodol-

Hydrothermal Reaction ogy is in the appraisal of exergonic reactions at
higher temperatures compared to endergonic
Koichiro Matsuno ones expected at lower temperatures, though it
Nagaoka University of Technology, Nagaoka, may look counterintuitive at first sight. A case in
Japan point is an estimate of the thermodynamic likeli-
hood of synthesizing amino acids from their pre-
cursors in moderately reducing hydrothermal
Keywords environments. Thermodynamic evaluation in
terms of the Gibbs free energy reveals that the
Exergonic reactions; Heating; Hot vents; Hydro- synthesis of many amino acids is exergonic when
lysis; Hydrothermal circulation; Quenching; synthesized from their component smaller mole-
Selective retention cules at temperature 100 C, whereas the synthe-
sis is endergonic at 18 C (Amend and Shock
1998).
Definition That synthesizing organic molecules such as
amino acids in hydrothermal environments is
Hydrothermal reactions are chemical reactions exergonic implies the synthetic reactions be
that take place in or near hydrothermal vents. Of energetically downhill and is quite welcome
importance from the perspective of prebiotic from the perspective of prebiotic synthesis at
chemistry is the fact that the two processes, least until finally reaching thermal equilibrium.
heating and quenching, are separated due to the Evolutionary relevance of chemical reactions in
repeated hydrothermal circulation of seawater thermal equilibrium is thus sought in how the
around the hot vents. Heating is indispensable initial reactants could be prepared and how the
for activating the reactants for a wide variety of reaction products could be removed. In fact,
synthetic reactions, while quenching is crucial for equilibrium prebiotic chemistry can provide a
selectively retaining some of those species that wide variety of reaction products depending
are synthesized. Heating and quenching experi- upon the choice of the initial reactants. Some
enced sequentially during hydrothermal circula- products might indeed catalyze reactions that
tion of seawater can make prebiotic chemistry have some prebiotic utility, but other products
evolutionary without being entrapped in thermal may well inhibit such catalysis or catalyze
equilibrium in which both heating and quenching adverse reactions. An example of this sort is
remain inseparable. seen in the clay-catalyzed condensation of
nucleotides, which could work best if only one
type of enantiomer of the starting material is
Overview present, while if both types were present, cata-
lytic condensation of one enantiomer could be
Hydrothermal reactions are classified into two disturbed and inhibited by the presence of the
distinct groups. One is for the reactions in thermal other enantiomer (Joyce and Orgel 1999).
equilibrium, and the other is for those off thermal Instrumental to making prebiotic chemistry
equilibrium. evolutionary must therefore be naturalization
Experimental examination of synthetic chem- of the coordination of both supplying the initial
ical reactions in thermal equilibrium starts from reactants to and removing the products from
the reactants supplied externally to an environ- reaction sites in an integrated manner. One nat-
ment initially prepared in thermal equilibrium ural candidate for meeting this challenge is con-
and specifies the character of the final synthetic stant circulation of seawater through the
products precipitated under the imposed equilib- periphery or the vicinity of hydrothermal vents,
rium conditions. An advantage of hydrothermal the latter of which could work as reaction sites
Hydrothermal Reaction 1161
because of the availability of heat energy for derivatives that could survive the preceding
activating various reactions. Both reactants and period of hydrolysis even partially may partici-
products are carried by the circulating flow of pate in the further elongation when they visit the
seawater and visit the hot reaction sites occa- hot spots the next time. The elongated products
sionally, while the flow stays in the vast cold are thus inclined to selectively maintain only
ocean during most of the time. Chemical reac- those oligomers of the kind that could be evolu-
tions proceeding within the circulating flow are tionary in keeping the preceding memory of syn-
undoubtedly not in thermal equilibrium due to thesis even partially.
crossing the temperature gradients generated There is nothing special with both thermal
around the vents repeatedly as dismissing the synthesis of products and their hydrolysis in pre-
occurrence of stationary chemical reactions biotic chemistry. However, if one conceives of
anchored at thermal equilibrium. both within the homogeneous scheme of chemi-
When one tries to figure out the average fre- cal reactions in thermal equilibrium, a formidable
quency of an arbitrary tiny droplet of seawater in obstacle would come up. No separation between
the ocean visiting one hydrothermal ridge sys- synthesis and hydrolysis is possible there. Syn-
tem, its modest estimate would give about once thesis and hydrolysis would produce a homoge- H
every 30 million years upon the conjecture that nized mixture in thermal equilibrium, with
the weight of the total seawater on the surface of ending up with a mere mixture of complex prod-
Earth is of the order of 1018 t and the water issued ucts and no chemical evolution.
from a single ridge system is about 1,000 t per What is specific to hydrothermal circulation of
second. If there are about 1,000 independent seawater around the hot vents in the ocean is its
hydrothermal ridge systems in the ocean even natural capacity of separating between synthetic
prior to the emergence of the major continents, production and selective retention of the prod-
the frequency of a tiny droplet visiting any one ucts. As a matter of fact, one may be able to
of the ridge systems could be raised to about observe how products would be synthesized in
once every 30 thousand years. This numerical the hot spots and how those products could sub-
exercise simply gives a possible lower limit to sequently survive their possible hydrolysis when
the actual visiting frequency. The actual visiting transferred into the cold surroundings by follow-
can however be made far more frequent locally ing the streamline of the circulating fluid sequen-
because of the participation of a wide variety of tially. What remains to be seen is to address the
shortcuts. tractable issue of whether the present picture of
Geology-assisted hydrothermal circulation of separating synthetic production from selective
seawater is peculiar in converting the preceding retention of the products could experimentally
reaction products in the solution into the be testable.
succeeding reactants when visiting hot spots One relevant experiment is the synthesis of
again and again, after being subject to rapid oligopeptides from monomeric glycine in a flow
quenching every time. This hydrothermal con- reactor simulating hydrothermal circulation of
version from products to reactants can have a seawater in the ocean, while setting the frequency
natural tendency to enhance the synthesis of of visiting a hot spot (225 C) to be once every
products of a specific kind. The underlying chem- 30–80 s instead of every 30,000 years (Imai
istry is pretty simple. Suppose one starts with a et al. 1999). When the frequency of visiting the
solution of monomers of a single kind to be hot spot was raised from once every 78 s to every
circulated. Elongation of oligomers out of the 34 s, the yields of oligomeric products were
initial monomers could be expected when both found to increase accordingly. The increase of
precursor oligomers and monomers are visiting the yields demonstrates the cumulative nature of
hot spots, while those products that could be synthetic production in the sense that the yields
hydrolyzed back into the original monomers are increased exponentially with the increase in the
released into the cold surroundings. Only those frequency of visiting the hot spot.
1162 Hydrothermal Vent Microbiology
Unless the synthetic production and selective See Also

retention of the products are forcibly homoge-
nized as in theoretical chemical reactions in ▶ Black Smoker, Organic Chemistry
thermal equilibrium, prebiotic chemistry pro- ▶ Hydrocarbons
ceeding in inhomogeneous thermodynamic con- ▶ Hydrothermal Vent Origin of Life Models
ditions (such as those applied to hydrothermal ▶ Polymer
circulation of seawater) could already be selec-
tive in retaining only a specific set of synthetic
products. Unique to the selective retention of the
products associated with the hydrothermal reac- Amend J, Shock EL (1998) Energetics of amino acid
tions is the built-in selective sieve for those synthesis in hydrothermal ecosystems. Science
products surviving rapid quenching that takes 281:1659–1662
place soon after the products leave the hot Imai E, Honda H, Hatori K, Brack A, Matsuno K (1999)
Elongation of oligopeptides in a simulated submarine
spots and enter into the cold environments. In hydrothermal system. Science 283:831–833
particular, there are at least two distinct fates Joyce GF, Orgel LE (1999) Prospects for understanding the
waiting for the product entering into the cold. origin of the RNA world. In: Guesteland RF, Cech TR,
One is to lower the temperature of the product Atkins JF (eds) The RNA world, 2nd edn. Cold Spring
Harbor Laboratory, Cold Spring Harbor, pp 49–77
with the aid of its hydrolysis, and the other is to Matsuno K, Nemoto A (2005) Quantum as a heat
lower the temperature of the product by storing engine – the physics of intensities unique to the origins
the acquired heat energy internally in the form of of life. Phys Life Rev 2:227–250
chemical bonds without suffering hydrolysis.
Which will win depends entirely upon which is
faster. This is the situation of winner takes all. Hydrothermal Vent Microbiology
Those that could not survive rapid quenching
would necessarily be pruned off. ▶ Hot Vent Microbiology
Prebiotic chemistry aimed at addressing the
origins of life has to face two different tasks in an
integrated manner at the same time. One task is
to prepare the chemistry that could be suffi- Hydrothermal Vent Origin of Life
ciently synthetic and versatile enough to be ame- Models
nable to evolutionary adaptation. And another
one is to secure the chemistry that could be Koichiro Matsuno and Eiichi Imai
sufficiently fine tuned and specific enough to Nagaoka University of Technology, Nagaoka,
maintain a well-orchestrated self-organization. Japan
At first sight, these two tasks would seem con-
tradictory. Despite that, chemical reactions rid-
ing on hydrothermal circulation of seawater may Keywords
suggest to us a loophole for reaching out to a
meaningful integration of the seemingly contra- Amino acids; Black smoker; Carboxylic acids;
dictory tasks by letting the circulating flow serve Concentration problem; Hydrothermal vent;
as a unifying thread. One likely means of exam- Membranous structures; Nucleotides; White
ining the present objective experimentally might smoker
be to try to run the overall exergonic reactions of
the oxidative citric acid cycle within the para-
digm of hydrothermal reactions under the pre- Definition
mise that pyruvate as the energy and carbon
source is already available (Matsuno and ▶ Hydrothermal environments are places where
Nemoto 2005). hot water comes into contact with mineral surfaces.
Hydrothermal Vent Origin of Life Models 1163
History It is likely that the geothermal heat flux was

greater near the time of the origin of life than at
Perhaps the first specific proposal (aside from present (Holm 1992); thus, hydrothermal sites
Darwin’s offhand reference to a warm little were likely more abundant on the early Earth,
pond (Calvin 1961), which is in some ways a and they may be more abundant in the modern
hydrothermal model) for the involvement of solar system, for example, in the interiors of Mars
hydrothermal environments in the ▶ origin of and Europa, than are favorable surface environ-
life was that of Harvey in 1924 (Harvey 1924), ments (CIBA 1996). Furthermore, it has also
who suggested that terrestrial thermal springs been suggested that hyperthermophilic
such as those found in Yellowstone National archaebacteria, of the sort typically associated
Park in Wyoming might have been ideal sites with submarine vents, cluster near the roots of
for chemical evolution. The idea was resurrected phylogenetic trees (Stetter 2006), supporting the
after the 1979 discovery of black smoker subma- idea that life originated at high temperature, or at
rine hydrothermal sites associated with least passed through an early high-temperature
mid-ocean spreading centers (Corliss bottleneck (Arrhenius et al. 1999).
et al. 1981). Since its modern resurrection, a There are a number of potential geochemical H
wealth of theoretical and experimental studies settings which are compatible with hydrothermal
have examined the idea from various perspec- origin of life models; among them are ▶ black
tives; nevertheless, the idea remains contentious. smoker type vents, off-axis type vents, and ter-
restrial hot spring environments. Black smoker
hydrothermal environments have typically very
Overview high temperature (350 C) and are acidic,
reducing, and sulfide rich. Significant amounts
The resurgence of the hydrothermal model in the of metal sulfides may be deposited in these envi-
late 1970s offered a counterpoint to the long- ronments. The eponymous black chimneys are
prevalent Oparin-Haldane heterotrophic model, largely composed of metal sulfides which precip-
which had attracted several criticisms. Among itate as the reducing, hot, acidic metal, and
them were the notion that the early atmosphere sulfide-rich hydrothermal fluids meet the cold
may not have been reducing (in which case the (2 C) oxidizing ambient seawater.
yield of organics from atmospheric synthesis Not long after the discovery of these sites,
would have been greatly reduced (Schlesinger Wächtershäuser (1988) proposed a pyrite surface
and Miller 1983)), the possibility that large mete- model for the origin of life, which suggested life
orite impacts might have repeatedly sterilized the arose based on the attachment of negatively
surface of the Earth (Maher and Stevenson 1988), charged molecules such as organophosphates
and the possible detrimental effects of UV radia- and carboxylic acids (which are ubiquitous in
tion on surface chemistry in the absence of an modern biochemistry) to positively charged
ozone layer (Berkner and Marshall 1965). pyrite surfaces. A series of carbon-fixing reac-
Among the proposed benefits of hydrothermal tions were proposed to be catalyzed by these
vent environments for the origin of life are thus surfaces which allowed for the duplication of
(1) protection from harsh surface conditions the catalytic surface reaction system, which sub-
(such as the aforementioned UV radiation and sequently spread across the mineral surface and
ocean sterilizing meteorite impacts); (2) an abun- complexified over time. This model was not ini-
dance of chemical potential in the form of tially explicitly proposed in the context of sub-
reduced dissolved gases and metals; (3) mineral marine hydrothermal vents, but the ubiquity of
surfaces to serve as catalysts, adsorbents, and sulfide minerals in such environments quickly led
cavities for the production and entrapment of to its being considered in that manner.
organic compounds; and (4) thermal energy to Debate soon focused on the extremely high
overcome kinetic barriers to reaction. temperatures and low pH values found in such
1164 Hydrothermal Vent Origin of Life Models
systems, conditions under which most organic systems, their reducing nature was not likely
compounds are rather unstable (White 1984), much greater than it is now. Presently, there is
which led many to doubt their appropriateness some debate over how much of the reduced gases
as sites for the accumulation and complexi- (which consist predominantly of H2, CH4, and
fication of organics. After this debate had sub- lesser amounts of ethane, ethane, propene, and
sided somewhat, a second type of submarine vent propane (Proskurowski et al. 2008)), and this
was discovered, the so-called Lost City or does appear to be in line with equilibration
peridotite-hosted hydrothermal vent system under the reducing environment of the system.
(Kelley et al. 2001). These environments have However, although the organic molecules
typically much lower temperature (40–90 C), detected thus far are extremely depleted in 14C,
and the fluid emanating from them is basic suggesting the source of the carbon in them is the
(pH 9–11) and quite reducing. Hydrothermal ori- mantle derived, the confirmation of the presence
gin of life models has arisen over time to accom- of more complex abiotic organic or reduced
modate the conditions more typically found in nitrogen compounds is frustrated by contaminat-
these environments (see, e.g., Martin and Russell ing biological material which becomes entrained
(2007)). as water is drawn into the vent system from
Laboratory simulations of the stability and neighboring pore sediment water. Thus, it
reactivity of organic compounds under hydro- remains uncertain whether even simple organics
thermal vent type conditions are complicated by can be generated in these systems. There is cer-
the fact that the conditions in these environments tainly room for further investigation of this
range so widely (i.e., from highly reducing to possibility.
highly oxidizing, from 2 to 350 C, from pH One suggested the advantage of synthesizing
2 to pH 11, and as fluids are in contact with organic compounds in the vicinity of hydrother-
diverse and variegated mineral surfaces for mal vents is that many synthetic reactions are
periods of time ranging from seconds to tens of calculated to be exergonic at elevated tempera-
thousands of years (Holm 1992)). tures and pressures. The syntheses of 11 of the
Some problems which need to be overcome in 20 protein-forming amino acids were calculated
hydrothermal vent origin of life models include to be exergonic at 100 C (Amend and Shock
concentrating reactants, synthesizing small 1998). However, Bada et al. (1995), among
organic molecules from dilute solutions under others, have shown that amino acids are
conditions that many organic molecules are extremely unstable at elevated temperature.
unstable, and the polymerization of those mono- Even simple molecules such as glycine have
mers into more complex molecules such as pep- short half-lives at temperatures above 150 C,
tides, oligonucleotides, and membranes (with the and the abundance of such molecules would
types of molecules specific to the particular depend on the balance between their synthesis
model of the proposer). and degradation. The experimental verification
While many have proposed that complex of such syntheses under conditions comparable
organic chemistry may occur in deep-sea hydro- to those found in natural environments is thus far
thermal vent environments, there are a number of lacking.
obstacles to this. First, the notion that the atmo- The synthesis of more complex molecules
sphere was not particularly reducing stems from such as nucleotides, oligonucleotides, lipids, or
the idea that the upper mantle had already peptides requires that several precursor mole-
released most of its volatiles and its reducing cules be brought together. As these reactions are
equivalents (largely in the form of metallic and at the very least kinetically second order, they
reduced iron) had already segregated into the require some concentration mechanism to occur
lower mantle and core (Delano 2001). Thus, appreciably. One proposed mechanism for solv-
although ▶ serpentinization may have been the ing the concentration problem is thermophoresis
source of reducing equivalents in hydrothermal in hydrothermal pore systems. Baaske
Hydrothermal Vent Origin of Life Models 1165
et al. (2007) have theoretically and experimen- as the bulk ocean would likely have had a very
tally discovered model laboratory systems that low concentration of organic compounds
suggest pore systems inside rocks in the vicinity (Aubrey et al. 2009). Terrestrial thermal pools
of ocean hydrothermal vents could enhance the may be more favorable in this regard.
concentration of reactants in pore water with the There are thus several problems for hydrother-
aid of the marked temperature gradients available mal vent models with respect to organic chemis-
there. The enhancement of the concentration is try. Nevertheless, several lines of evidence point
due to the interplay between convective and dif- toward an early high-temperature period for life,
fusive flow of solutions in the presence of tem- and there also remain numerous unknown aspects
perature gradients. Another proposed solution is of the types of chemistry which might occur in
surface adsorption; however, mineral surfaces are such environments which warrant further study.
also known to act as catalysts for breaking molec-
ular bonds (Marshall-Bowman et al. 2010), and
surface adsorption typically becomes weaker at
high temperature; thus, most organics would See Also
likely tend to migrate to lower-temperature envi- H
ronments (Cohn et al. 2001), and the residence ▶ Amino Acid
time of molecules in the high-temperature por- ▶ Black Smoker
tion of the systems also requires some experimen- ▶ Black Smoker, Organic Chemistry
tal verification. ▶ Earth, Formation and Early Evolution
Provided significant starting concentrations of ▶ Hot Vent Microbiology
amino acids, peptides can be formed at elevated ▶ Hydrothermal Environments
temperature in water, both with and without con- ▶ Hydrothermal Reaction
densing agents, as long as the reactions are not ▶ Oligopeptide
heated for very long. For example, tripeptides of ▶ Origin of Life
phenylalanine, tyrosine, and glycine have been ▶ Polymer
synthesized at 100 C in slurries of FeS and NiS ▶ Serpentinization
as a catalyst in the presence of hydrogen sulfide ▶ White Smoker
or methanethiol as a condensing agent (Huber
and Wächtershäuser 1998). However, peptides References and Further Reading
are thermally unstable in water. Below mM con-
centrations of amino acids, peptides do not form Amend JP, Shock EL (1998) Energetics of amino acid
appreciably (Cleaves et al. 2009). Peptides are synthesis in hydrothermal ecosystems. Science
281:1659–1662
considerably more stable than oligonucleotides; Arrhenius G, Bada JL, Joyce GF, Lazcano A, Miller S,
thus, these sorts of considerations are likely more Orgel LE (1999) Origin and ancestor: separate envi-
severe for molecules such as RNA, presenting a ronments. Science 283:792
disconnect between hydrothermal vent models Aubrey AD, Cleaves HJ, Bada JL (2009) The role of
submarine hydrothermal systems in the synthesis of
and RNA World models. amino acids. Orig Life Evol Biosph 39:91–108
Cycling could overcome some of these prob- Baaske P, Weinert FM, Duhr S, Lemke KH, Russell MJ,
lems. For example, molecules could be synthe- Braun D (2007) Extreme accumulation of nucleotides
sized during residence in a high-temperature in simulated hydrothermal pore systems. Proc Natl
Acad Sci U S A 104:9346–9351
milieu and then quickly quenched into a lower- Bada JL, Miller SL, Zhao M (1995) The stability of amino
temperature region. Imai et al. (1999) observed acids at submarine hydrothermal vent temperatures.
that the synthesis of hexamer of glycine was Orig Life Evol Biosph 25:111–118
possible from the cyclic reaction of fairly con- Berkner LV, Marshall LC (1965) On the origin and rise of
oxygen concentration in the earth’s atmosphere.
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deep-sea environments, this may be problematic, Ann Intern Med 54:954–976
1166 Hydroxy Acid
Cleaves H, Aubrey A, Bada J (2009) An evaluation of the

critical parameters for abiotic peptide synthesis in Hydroxy Acid
submarine hydrothermal systems. Orig Life
Cohn CA, Hansson TK, Larsson HS, Sowerby SJ, Holm Henderson James (Jim) Cleaves II
NG (2001) Fate of prebiotic adenine. Astrobiology Earth–Life Science Institute (ELSI), Tokyo
1(4):477–480 Institute of Technology, Meguro–ku, Tokyo,
Corliss JB, Baross JA, Hoffman SE (1981) An hypothesis
concerning the relationship between submarine hot Japan
springs and the origin of life on Earth. Oceanol Acta Institute for Advanced Study, Princeton, NJ,
4:59–69 USA
Delano JW (2001) Redox history of the Earth’s interior Blue Marble Space Institute of Science,
since approximately 3900 Ma: implications for prebi-
otic molecules. Orig Life Evol Biosph 31:311–341 Washington, DC, USA
Foundation Symposium CIBA (1996) Evolution of hydro- Center for Chemical Evolution, Georgia Institute
thermal ecosystems on Earth (and Mars) (Novartis of Technology, Atlanta, GA, USA
Foundation Symposia). Wiley, Chichester
Harvey RB (1924) Enzymes of thermal algae. Science
60:481–482
Holm NG (1992) Marine hydrothermal systems and the
origin of life: report of SCOR Working Group 91. Definition
Springer, Berlin
Huber C, Wächtershäuser G (1998) Peptides by activation
of amino acids with CO on (Ni, Fe)S surfaces: impli- Hydroxy acids are organic compounds containing
cations for the origin of life. Science 281:670–672 both hydroxyl and carboxylic acid functional
Imai E, Honda H, Hatori K, Brack A, Matsuno K (1999) groups. a-Hydroxy acids are a particularly
Elongation of oligopeptides in a simulated submarine important class of carboxylic acids which
hydrothermal system. Science 283:831–833
Kelley DS, Karson JA, Blackman DK, Fr€ uh-Green G, have a hydroxyl group on the carbon adjacent
Gee J, Butterfield DA, Lilley MD, Olson EJ, Roe to the carboxylic acid group or a-carbon.
KR, Shipboard Scientific Party (2001) An off-axis They may be either naturally occurring
hydrothermal vent field near the Mid-Atlantic Ridge or synthetic. Commonly occurring
at 30 N. Nature 412:145–149
Maher KA, Stevenson DJ (1988) Impact frustration of the biological a-hydroxy acids include
origin of life. Nature 331:612–614 lactic (CH3CHOHCOOH), glyceric
Marshall-Bowman K, Ohara S, Sverjensky DA, (HOCH2CHOHCOOH), glycolic (HOCH2COOH),
Hazen RM, Cleaves HJ (2010) Catalytic peptide and malic (HOOCCH2CHOHCOOH) acids.
hydrolysis by mineral surface: implications for prebi-
otic chemistry. Geochim Cosmochim Acta 74: a-Hydroxy acids are important metabolic products
5825–5861 of the biological transamination of ▶ amino
Martin W, Russell MJ (2007) On the origin of biochemis- acids. Several a-hydroxy acids and a-hydroxy
try at an alkaline hydrothermal vent. Philos Trans acid polyols have been identified in the Murchison
R Soc Lond B 362:1887–1925
Proskurowski G, Lilley MD, Seewald JS, Fr€ uh-Green GL, meteorite. Some interpret the presence of
Olson EJ, Lupton JE, Sylva JP, Kelley DS (2008) Abio- a-hydroxy acids in this body as evidence for the
genic hydrocarbon production at lost city hydrother- Strecker-cyanohydrin mechanism for the synthesis
mal field. Science 319:604–607 of the amino acids also found in it.
Schlesinger G, Miller S (1983) Prebiotic synthesis in
atmospheres containing CH4, CO, and CO2. J Mol
Evol 19:383–390
Stetter KO (2006) Hyperthermophiles in the history of
life. Philos Trans R Soc B 361:1837–1843 See Also
Wächtershäuser G (1988) Before enzymes and templates:
theory of surface metabolism. Microbiol Rev ▶ Amino Acid
52(4):452–484
▶ Carboxylic Acid
White RH (1984) Hydrolytic stability of biomolecules at
high temperatures and its implication for life at 250 - ▶ Hydroxyl Group
C. Nature 310:430–432 ▶ Strecker Synthesis
Hydroxyl Radical 1167
Hydroxyacetaldehyde Hydroxyl Radical
▶ Glycolaldehyde William M. Irvine

2-Hydroxyethanoic Acid
Synonyms
▶ Glycolic Acid
OH
Hydroxyl Group
Definition
Earth–Life Science Institute (ELSI), Tokyo The hydroxyl radical, containing single oxygen H
Institute of Technology, Meguro–ku, Tokyo, and hydrogen atoms, is highly reactive. In the
Japan terrestrial troposphere, it interacts with and
Institute for Advanced Study, Princeton, NJ, removes a number of pollutants as well as the
USA greenhouse gases methane and ozone. It can
Blue Marble Space Institute of Science, damage typical biological macromolecules.
Washington, DC, USA As a free diatomic molecule, it is widespread in
Center for Chemical Evolution, Georgia Institute the ▶ interstellar medium of our Milky
of Technology, Atlanta, GA, USA Way galaxy and other galaxies, where it is fre-
quently observed in maser emission. Some
external galaxies contain so-called OH
Definition megamasers, with luminosity orders of magni-
tude greater than masers in our Galaxy. In the
The term hydroxyl group is used to describe the solar system, OH is a major constituent of com-
-OH functional group in an organic compound. etary comae and an indirect measure of their
Organic molecules containing hydroxyl groups water content.
are known as alcohols. Hydroxyl groups are espe-
cially important in biochemistry due to their ten-
dency to form hydrogen bonds, as they contain both
lone pairs of electrons and a weakly acidic proton.
History
Hydroxyl groups are also able to form hydrogen
The OH radical was the first interstellar molecule
bonds with water, a property that increases the
detected by radio astronomers, in 1963 by
hydrophilicity and solubility of molecules
Weinreb et al. Measurements of the Zeeman
containing them. The carbohydrates are an exam-
effect for OH transitions have subsequently
ple of a group of molecules that are extremely
been used to estimate the magnetic field in the
soluble due to hydroxyl functional groups.
interstellar medium. In 1981, the pure rotational
lines of OH were observed at far-infrared wave-
See Also lengths in absorption against the continuum emis-
sion from a background molecular cloud.
▶ Alcohol Cometary OH has been detected both in the ultra-
▶ Carbohydrate violet and, in 1973, at radio wavelengths; this led
1168 4-Hydroxyphenylalanine
to the correct prediction that H2O ice was present

in cometary nuclei and was the source of the Hygiea
hydroxyl radical (through photodissociation by
the solar ultraviolet photons). Carmen Tornow
Institute of Planetary Research, German
Aerospace Center, Berlin, Germany
See Also
Definition
▶ Comet (10) Hygiea, named after the Greek goddess of
▶ Interstellar Medium health, is located in the main asteroid belt
▶ Maser between 2.77 and 3.50 AU (http://ssd.jpl.nasa.
▶ Molecules in Space gov/sbdb.cgi). According to its inclination and
eccentricity it belongs to the core population of
the main belt involving more than 90 % of the
References and Further Reading asteroids. Hygiea revolves in 5.56 years around
the sun and rotates in 27.6 h around itself. With a
Bockelee-Morvan D, Crovisier J, Mumma MJ, Weaver radiometric diameter between 407 and 469 km
HA (2004) The composition of cometary volatiles. and with a density of 2,080 kg/m3, it is the fourth
largest asteroid by volume and mass after
II. The University of Arizona Press, Tucson,
pp 391–423 (1) Ceres, (4) Vesta, and (2) Pallas. Hygiea’s
Hartquist TW, Williams DA (1995) The chemically con- shape corresponds to a very oblate spheroid,
trolled cosmos. Cambridge University Press, while the three larger asteroids have clearly
Cambridge
Weinreb S, Barrett AH, Meeks L, Henry JC (1963) Radio
less oblate shapes but comparatively short
observations of OH in the interstellar medium. Nature rotation periods below 10 h. As a member of
200:829–831 the large population of C-type asteroids,
Hygiea’s chemical surface composition pro-
duces a spectral reflectance signature which
resembles the one of carbonaceous chondrites
subjected to thermal processing and aqueous
4-Hydroxyphenylalanine alteration.
▶ Tyrosine
History
Despite its large size, Hygiea was discovered

2-Hydroxypropanoic acid relatively late by Annibale de Gasparis on April
12, 1849. The detection of the rather large aster-
▶ Lactic Acid oid has taken a long time compared to (2) Pallas
or (3) Juno, which were observed in 1802 and
1804, because of Hygiea’s low geometric albedo
and rather large semi-major axis. As a result the
asteroid has an apparent magnitude between
6-Hydroxypurine 9 and 12. Accordingly it is no surprise that
Hygiea’s asteroid family was detected not until
▶ Hypoxanthine 1978 by Andrea Carusi and Enrico Massaro.
Hypercycle 1169
Today the existence of this family is verified by a

1
number of independent observations.
See Also n 2
▶ Asteroid
▶ Ceres
▶ Juno
4 3
▶ Orbit
▶ Pallas
Hypercycle, Fig. 1 The catalytic interactions in a
▶ Taxonomy hypercycle with n members. The individual elements,
▶ Vesta 1, 2, 3, 4, . . ., n, are coupled by cyclic catalysis (black H
arrows): 1!2, 2!3, 3!4, 4!n, n!1. Every member is
an autocatalyst as indicated by cycles (small circular
arrows)
Hypercycle History
Peter Schuster The hypercycle was invented in the 1960s (Eigen

Institut f€
ur Theoretische Chemie der Universität 1971) as a model for cooperation of otherwise
Wien, Wien, Austria competing elements that are capable of self-
replication. Hypercycles were postulated as
mechanisms for major transitions in evolution
Keywords (Eigen and Schuster 1977, 1978a, b; Maynard
Smith and Szathmáry 1995; Schuster 1996;
Autocatalytic network; Hyperbolic growth; Phillipson and Schuster 2009).
Major transitions; Dynamical systems; Oscilla-
tions; Symbiosis
Overview
Several replicators (X1, X2, . . ., Xn) – entities

Synonyms
capable of self-replication – form a cyclic cata-
lytic network in which the replication is catalyzed
Cyclic replicator equation
by the precursor element: X1 catalyzes the forma-
tion of X2 and so on until Xn catalyzes the forma-
tion of X1. Thereby, the catalytic interactions
Definition form a closed loop: 1!2!3!. . .!n!1. The
simplest theoretical example is the elementary
A hypercycle is a catalytic network of hypercycle, which in mathematical terms is
autocatalysts (Ik) that are functionally coupled described by the differential equation
in a dynamical cycle: ) I1 ) I2 ) . . .
dxj Xn
) In ) I1 ) . Hypercycle dynamics are ¼ xj fj xj1 fx x ;
dt i¼1 i i1 i
described by means of nonlinear differential equa- (1)
tions of replicator equation type (Fig. 1). i, j ¼ 1, 2, . . . , n and i, j ¼ mod n
1170 Hypercycle
Hypercycle, Fig. 2 Hypercycle dynamics visualized in damped oscillations; n = 4, the concentrations approach
form of solution curves of Eq. 1. Normalized concentra- the stationary values with weakly damped oscillations;
tions of the members of hypercycles xj(t) are plotted as and n = 5, the concentrations oscillate without damping.
functions of time t for different sizes of the cycle. Indi- For n > 5, hypercycle dynamics shows oscillations with-
vidual members (different colors) cooperate in the repro- out damping as illustrated here for the n = 5 case. The
duction of the entire cycle. Different dynamics is observed following parameter values were chosen: f1 = f2 = f3 =
for different sizes: n = 2, the concentrations approach f4 = f5 = 1; initial conditions, x1 ð0Þ ¼ 0:95 and
their stationary values monotonously; n = 3, the concen- x2 ð0Þ ¼ 0:05 for n = 2 and x1 ð0Þ ¼ 0:9 and
trations approach the stationary values with strongly xk ð0Þ ¼ 0:1=ðn 1Þ for n = 3, 4, 5
which is a special case of replicator equations curves xj (t) show characteristic dependence on
(Hofbauer and Sigmund 1998; Nowak 2006). the size of the hypercycle (Fig. 2):
Xn
The variables are
normalized concentrations of
replicators,
Xn X j ¼ N j and x j ¼ N j = N
i¼1 i 1. For n = 2, the dynamical system converges
with x
j¼1 j
¼ 1 . The parameters fj are rate monotonously to a unique and asymptotically
parameters of the catalyzed replication reactions, stationary state at which both replicators X1
ðAÞ þ Xj þ Xj1 ! 2Xj þ Xj1, where (A) stands and X2 are present.
for the building blocks for the synthesis. 2. For n = 3 and n = 4, the stationary state is
The qualitative features of solutions of Eq. 1 unique and asymptotically stable. The solution
do not depend on particular values fj curves xj (t) show strongly damped oscilla-
(as long as they are strictly positive, fj > 0, tions n = 3 and weak damping for n = 4.
Hofbauer and Sigmund 1998). Without losing 3. For n 5, the stationary state is unstable and
generality, all fj values are chosen to be equal, the concentrations xj (t) oscillate.
and the dynamical system has a stationary
point in the middle of concentration space: Independently of internal dynamics, the total
x1 ¼ x2 ¼ x3 ¼ . . . ¼ xn ¼ 1=n . The solution concentrations of hypercycles as integral units,
Hypersaline Lakes 1171
Xn
CðtÞ ¼ N ðtÞ, grow stronger than exponen-
i¼1 i
Phillipson PE, Schuster P (2009) Modeling by nonlinear
differential equations. Dissipative and conservative
tial for unlimited resources. This hyperbolic processes. World Scientific, Singapore, pp 61–75
growth is observed for a class of systems in Schuster P (1996) How does complexity arise in evolu-
which the population size has an enhancing effect tion? Complexity 2(1):22–30
Vaidya N, Manapat ML, Chen IA, Xulvi-Brunet R,
on the growth rate. Examples of hyperbolic growth Hayden EJ, Lehman N (2012) Spontaneous network
were observed in virology, in demography (growth formation among cooperative RNA replicators. Nature
of human population), and in economics 491:72–77
(increasing returns). Systems with hyperbolic
growth outgrow exponentially growing systems,
since in theory a hyperbolically growing popula-
tion can reach infinity in finite times. Once a Hypersaline Environment
hypercycle has been established, it is very hard to
replace it by another system: hypercycles are can- Felipe Gomez
didates for once-and-forever decisions. Examples Centro de Astrobiologı́a (CSIC/INTA), Instituto
for hypercycles in the real world are symbioses Nacional de Técnica Aeroespacial, Torrejón de
Ardoz, Madrid, Spain H
and other forms of cooperation within and between
species. Formation of hypercycle-like networks
among cooperative RNA replicators has been
reported (Vaidya et al. 2012). Definition
Hypersaline environment is an extreme habitat

See Also dominated by high salt concentration. Associ-
ated to this extreme condition, other extreme
▶ Endosymbiosis conditions can be found like low oxygen con-
▶ Evolution, Biological centration and alkaline conditions. The inhabi-
▶ Evolution, Molecular tants of these environments are microorganisms
▶ Quasispecies that are called “halophiles.” Halophiles have to
▶ Self-Replication solve the problem of the osmotic pressure gen-
▶ Symbiosis erated in the cell membrane. To do that, they can
use compatible solutes. Different groups can be
References and Further Reading distinguished on the basis of their physiological
responses to salt. Several classifications have
Eigen M (1971) Self-organization of matter and the evo- been proposed; one considers the optimum
lution of biological macromolecules. Naturwis-
senschaften 58:465–523
growth of the microorganisms at different salt
Eigen M, Schuster P (1977) The hypercycle. A principle concentrations: non-halophiles are those that
of natural self-organization. Part A: emergence of the grow best in media containing less than 0.2 M
hypercycle. Naturwissenschaften 64:541–565 NaCl, halotolerant microorganisms that can
Eigen M, Schuster P (1978a) The hypercycle. A principle
of natural self-organization. Part B: the abstract
tolerate high salt concentrations, moderate
hypercycle. Naturwissenschaften 65:7–41 halophiles growing between 0.2 and 2.5 M
Eigen M, Schuster P (1978b) The hypercycle. A principle NaCl, and extreme halophiles growing in opti-
of natural self-organization. Part C: the realistic mal conditions at high salt concentration
hypercycle. Naturwissenschaften 64:341–369
Hofbauer J, Sigmund K (1998) Evolutionary games and
(2.5–5.2 M NaCl).
replicator dynamics. Cambridge University Press,
Cambridge
Maynard Smith J, Szathmáry E (1995) The major transi-
tions in evolution. WH Freeman, Oxford
Nowak MA (2006) Evolutionary dynamics. The Belknap
Hypersaline Lakes
Press of Harvard University Press, Cambridge, MA,
Exploring the equations of life ▶ Soda Lakes
1172 Hypertelescope
Karl O. Stetter and Wolfram Zillig (Zillig

Hypertelescope et al. 1981; Stetter et al. 1981). In the following
two decades, further work led mainly by
▶ Interferometry K. O. Stetter focused on submarine or subterra-
nean thermal environments. This work allowed
the isolation and description of more than 50 spe-
cies of hyperthermophilic Archaea and a few
Bacteria, some requiring temperatures above
Hyperthermophile 90 C to grow, with optimum temperatures
above 100 C. Phylogenetic analysis based on
Jose Berenguer 16S RNA revealed that all of them are slowly
Centro de Biologı́a Molecular Severo Ochoa, evolving groups, supporting an ancient origin for
UAM-CSIC, Madrid, Spain this type of microorganisms.
Keywords Overview
High temperature; Hot springs; Hydrothermal Since the first descriptions of hyperthermophiles
vents; Origin of life; Thermal environment in 1981 (Zillig et al. 1981; Stetter et al. 1981),
about 90 species of hyperthermophilic Archaea
and Bacteria have been described from
Definition different terrestrial and marine thermal environ-
ments. Taxonomically and despite the existence
The word hyperthermophile (literally extremely of yet unassigned isolates, most are grouped in
heat loving) refers to a microorganism that has an ten orders (Archaea: Thermoproteales, Desulfur-
optimum temperature for growth above ococcales, Pyrodictiales, Thermococcales,
80 C. Most hyperthermophiles belong to the Archaeoglobales, Methanococcales,
Archaea domain, with only few exceptions Methanobacteriales, Methanopyrales; Bacteria,
belonging to Bacteria. Aquificales and Thermotogales). A few hyper-
thermophilic isolates are even unable to grow
below 80–90 C. Despite this, it has been possible
History to isolate hyperthermophiles far from the nearest
known thermal effluent, probably because they
Thomas D. Brock (1978) isolated the archaeal remain viable for years at low temperatures. At
genus Sulfolobus at Yellowstone National Park the upper part of the temperature range, there are
and showed that it grew optimally at 75 C with hyperthermophiles that can tolerate temperatures
an upper temperature limit for growth at 85 C. normally used in sterilization processes (121 C)
Previous reports by the same author described for 1 h (Pyrodictium occultum). Kashefi and
prokaryotic microorganisms that were able to Lovley (2003) reported residual growth of a
grow on microscopy slides inside boiling hyperthermophilic isolate under such conditions.
(92 C) hot springs. These organisms probably The use of different phylogenetic markers
were true hyperthermophiles. However, the first assigns the hyperthermophiles to the root of the
hyperthermophiles isolated and characterized in corresponding evolutionary trees, leading to the
the laboratory (Methanothermus fervidus and hypothesis that this kind of microorganism is
the Thermoproteus spp.) were isolated from very ancient. Despite this generally accepted
hot springs in Iceland and described simulta- view of the phylogenetic tree, the slow-evolving
neously in 1981 in two articles coauthored by character of hyperthermophiles has been
Hyperthermophile 1173
challenged by different studies in the last decade Carbon assimilation by hyperthermophiles grow-
based on other phylogenetic approaches ing chemolithotrophically takes place on CO2 or
(Ciccarelli et al. 2006). According to this view, CO mainly through the reductive acetyl coenzyme
hyperthermophilic character could be an adapta- A cycle (acetyl-CoA or Wood-Ljungdahl path-
tion of moderate ▶ thermophiles to superheated way), like in many anaerobic Gram-positive bac-
environments, in some cases through lateral gene teria, or through the reductive tricarboxylic acid
transfer. cycle (rTCA or Arnon cycle), as in green sulfur
Adaptation to hyperthermophilic conditions is bacteria, although alternative modified pathways
the result of a series of specific modifications of also exist. These cycles usually generate acetyl
the structures and macromolecules of the cell, as coenzyme A, from which gluconeogenesis must
simple metabolites and cofactors are basically the start. The presence of an irreversible condensing
same as those of mesophiles. Membrane adapta- fructose bisphosphate aldolase/phosphatase is a
tion results from an increase in the hydrophobic generalized trait within autotrophic hyperther-
lateral interactions between the major lipids, mophiles that appears to be an essential step to
esters of glycerol and two fatty acids in Bacteria, guide these pathways toward gluconeogenesis
and ethers of glycerol and two isoprenol-derived (Say and Fuchs 2010). Facultative and obligate H
alcohols in Archaea. In hyperthermophilic chemoorganotrophs also exist among hyperther-
Archaea, bipolar tetraether lipids form membrane mophiles that grow by anaerobic respiration,
monolayers in which C40 isoprenols are fre- essentially with elemental sulfur (S ) as electron
quently cyclized as a further thermal adaptation. acceptors or, more rarely, by fermentation (i.e.,
A low DNA content (1.8–3 Mbp) and the pres- Thermoproteus uzonensis). Data obtained from
ence of histone-like proteins have been described the analysis of thermophilic environments using
in some hyperthermophilic Archaea. The pres- molecular ecology methods (Pagé et al. 2008)
ence of type I topoisomerase that introduces pos- suggests that the biodiversity of chemoorga-
itive coils (reverse gyrase) into DNA is the only notrophic hyperthermophiles is underestimated.
trait common to hyperthermophilic Bacteria and
Archaea, leading to the hypothesis that DNA
stability is an important factor for life at high Basic Methodology
temperatures. However, Thermococcus
kodakaraensis mutants lacking this activity The high sensitivity to oxygen of hyperther-
show only a slight decrease in growth rate mophiles at their growth temperatures is an
above 90 C, so it has been proposed that this important challenge for their isolation. Environ-
enzyme could help to rewind ssDNA regions mental samples to be used for the isolation of
separated by thermal stress (Sandman 2008). hyperthermophiles (water, soil, rock, etc.) have
Enzymes from hyperthermophiles usually have to be taken under reducing conditions to prevent
an activity profile that fits the temperature growth any contact of microorganisms with oxygen at
range of the organism of origin as the result of a least at high temperature, when its toxic effect is
higher number of intra- and intermolecular inter- more severe. For terrestrial surface environ-
actions (Vieille and Zeikus 2001). ments, taking samples does not require sophisti-
Most hyperthermophiles grow as obligate or cated equipment. In the absence of N2 gas,
facultative chemolithotrophs by using either reductants such as sodium sulfide/sodium
hydrogen or sulfides (e.g., pyrite) as electron dithionite and an appropriate redox indicator
donors and nitrate, CO2, sulfur, sulfate, or ferric such as resazurin can be added to sample glass
iron as electron acceptors to gain energy. Low containers, which have to be tightly closed
concentrations of oxygen are also used by some with gas-impermeable butyl rubber stoppers.
isolates, but for most hyperthermophiles oxygen is Samples can then be transported to the labora-
highly toxic at or near their temperature of growth. tory at room temperature. For undersea hot
1174 Hyperthermophile
environments, perforation drills, water sam- (Blöchl et al. 1997), whereas description of
plers, piston corers, grab samplers, and dredge an isolate able to grow at 121 C is still under
samplers or even deep submergence vehicles discussion (Kashefi and Lovley 2003). A further
(DSVs) and remotely operative vehicles area of research on hyperthermophiles is focused
(ROVs) are required. Numbers of whole micro- on the impressive thermostability of their
bial communities in different submarine hydro- enzymes and their possible use in industrial pro-
thermal samples range from 104 to 109 cells per cesses. This thermostability is also responsible
gram at the surface of ▶ black smoker chimneys, for the ease with which their proteins and mac-
sediments, and hydrothermal plumes. Super- romolecular complexes crystallize under labora-
heated vent fluids can range from undetectable tory conditions, raising additional interest that
numbers up to 106 cells per ml (Nakagawa and led to develop large structural genomics projects
Takai 2006). (Jenney and Adams 2008).
Once at the laboratory, isolation of new
hyperthermophiles requires enrichment cultures
subjected to conditions mimicking those of the Applications
environment at the sampling site regarding tem-
perature, putative electron donors and acceptors, The high stability of enzymes under high temper-
and carbon sources. Growth rate is a limiting atures (there are enzymes active above 125 C) is
factor for successful enrichment, and periods of associated with an increase in compactness that
days are usually required before attempting the results in increased resistance to additional stresses
isolation of individual cells. Due to obvious dif- such as the presence in the reaction of detergents or
ficulties in keeping solid media under the pre- organic solvents (Vieille and Zeikus 2001). Such
scribed conditions, most isolation procedures combination of resistances fits specific applica-
require liquid medium and use either dilution tions in different fields. For example, DNA poly-
or physical methods such as optical tweezers. merases from hyperthermophiles (Pyrococcus sp.,
For high scale production, special fermenters Thermococcus sp.) are commonly used to amplify
have been designed to withstand the harsh DNA sequences with high fidelity. Other enzymes
growth conditions that most hyperthermophiles from hyperthermophiles are or have the potential
require, which in some case combine high tem- to be used in industrial processes for which high
peratures and pressure in acidic media (Stetter temperatures represent an added value, such as
2006). starch and cellulose hydrolysis and further sugar
modification (endoglucanases, glucosidases,
xylanases, amylases, glucoamylases, pullulanases,
Key Research Findings etc.), proteases for detergents, lipases and esterases
for biocatalysis, oxidoreductases like alcohol or
The main interest of hyperthermophiles is based glutamate dehydrogenases for biotransformations,
on two key observations. On the one hand, they C-C bonding enzymes (aldolases, transketolases),
were and still are regarded as ancestral organ- nitrile-degrading enzymes, etc. (Antranikian
isms, giving an opportunity to analyze what are 2008). Additional applications in specific but
most likely “living fossils” from the Archaean robust biosensor devices or for nanotechnology
era. This view is still the most accepted hypoth- are also in development. In addition to the
esis, although an alternative hypothesis has been enzymes, hyperthermophiles produce special
proposed. On the other hand, a search for the kinds of compatible solutes such as mannosyl
temperature limits for life has driven the glycerate, cyclic 2,3-biphosphoglycerate, and
search quest for more thermophilic microorgan- diglyceryl phosphate in relevant amounts, some
isms. Today, the upper temperature limit for of which have been shown to confer
growth of a well-established and described thermoprotection and protection against desicca-
microorganism is 113 C for Pyrolobus fumarii tion to mesophilic enzymes (Santos et al. 2008).
Hyperthermophile 1175
Future Directions References and Further Reading
Given the biological and applied interest of Antranikian G (2008) DNA-binding proteins and DNA
topology. In: Robb F, Antranikian G, Grogan D,
hyperthermophiles, future research in the field
Driessen A (eds) Thermophiles, biology and technol-
will develop several lines of interest. Their ancient ogy at high temperatures. CRC, Boca Raton,
nature and extreme form of living require a com- pp 113–160
prehensive physiological study of at least a pair of Blöchl E, Rachel R, Burggraf S, Hafenbradl D, Jannasch
HW, Stetter KO (1997) Pyrolobus fumarii, gen. and
model organisms, for which a reliable system for
sp. nov., represents a novel group of archaea,
genetic manipulation should be developed. Present extending the upper temperature limit for life to 113 -
developments on Thermococcus kodakaraensis C. Extremophiles 1:14–21
and Sulfolobus solfataricus are at the front lines Brock TD (1978) Thermophilic microorganisms and life
at high temperatures. Springer, Berlin
of this effort. The construction of efficient systems
Ciccarelli FD, Doerks T, von Mering C, Creevey CJ,
for the overexpression of genes in such models will Snel B, Bork P (2006) Toward automatic reconstruc-
widen the number of hyperthermophilic enzymes tion of a highly resolved tree of life. Science
available for applications that at present are hidden 311(5765):1283–1287
because of the inability to produce them in
Jenney FE Jr, Adams MW (2008) The impact of H
extremophiles on structural genomics (and vice
mesophilic hosts. Such expression systems will versa). Extremophiles 12:39–50
also be required to crystallize some of these pro- Kashefi K, Lovley DR (2003) Extending the upper tem-
teins and complexes. Also, the metagenomic perature limit for life. Science 301:934
Nakagawa S, Takai K (2006) The isolation of thermo-
approaches on hyperthermophilic environments
philes from deep-sea hydrothermal environments. In:
will open up a new world of enzymes having Rainey FA, Oren A (eds) Extremophiles. Methods in
biotechnological potential to help to optimize microbiology, vol 35. Elsevier, Amsterdam, pp 55–91
existing processes and develop new ones. Finally, Pagé A, Tivey MK, Stakes DS, Reysenbach AL (2008)
Temporal and spatial archaeal colonization of hydro-
molecular ecology methods in such environments
thermal vent deposits. Environ Microbiol 10:874–884
will likely unveil the presence of new Sandman K (2008) DNA-binding proteins and DNA
non-cultivable hyperthermophiles that could topology. In: Robb F, Antranikian G, Grogan D,
uncover some phylogenetic surprises. Hyperther- Driessen A (eds) Thermophiles, biology and technol-
ogy at high temperatures. CRC, Boca Raton,
mophiles are of special interest for astrobiology
pp 279–289
because high-temperature processes are important Santos H, Lamosa P, Faria TQ, Pais TM, Lopez de la
factors in the development of planetary bodies; Paz M, Serrano L (2008) DNA-binding proteins and
thus, the scenarios in which life can develop in DNA topology. In: Robb F, Antranikian G, Grogan D,
Driessen A (eds) Thermophiles, biology and technol-
the universe increase correspondingly.
ogy at high temperatures. CRC, Boca Raton, pp 9–24
Say RF, Fuchs G (2010) Fructose 1, 6-bisphosphate aldol-
ase/phosphatase may be an ancestral gluconeogenic
See Also enzyme. Nature 464:1077–1081
Stetter KO (2006) History of discovery of the first
▶ Autotroph hyperthermophiles. Extremophiles 10:357–362
Stetter KO, Thomm M, Winter J, Wildgruber G, Huber H,
▶ Autotrophy
Zillig W, Janecovic D, König H, Palm P, Wunderl
▶ Black Smoker S (1981) Methanothermus fervidus, sp. nov., a novel
▶ Chemolithoautotroph extremely thermophilic methanogen isolated from an
▶ Deep-Sea Microbiology Icelandic hot spring. Zbl Bakt Hyg I Abt Orig
C2:166–178
▶ Deep Subsurface Microbiology
Vieille C, Zeikus GJ (2001) Hyperthermophilic enzymes:
▶ Hot Spring Microbiology sources, uses, and molecular mechanisms for thermo-
▶ Hot Vent Microbiology stability. Microbiol Mol Biol Rev 65:1–43
▶ Hydrothermal Environments Zillig W, Stetter KO, Schäfer W, Janekovic D, Wunderl S,
Holz I, Palm P (1981) Thermoproteales: a novel type
of extremely thermoacidophilic anaerobic
▶ Osmolite archaebacteria isolated from Icelandic solfataras. Zbl
▶ Thermophile Bakt Hyg I Abt Orig C2:205–227
1176 Hypolithic
colonization of quartz stones has now been

Hypolithic documented in many locations, for example, the
Vestfold Hills, ▶ Antarctica (Smith et al. 2000);
Charles S. Cockell the Mojave Desert, USA (Schlesinger
Geomicrobiology Research Group, PSSRI, Open et al. 2003); the Negev desert, Israel (Berner
University, Milton Keynes, UK and Evenari 1978); the Namib Desert, Africa
(Budel and Wessels 1991); and the deserts of
China (Warren-Rhodes et al. 2007). On the
Keywords underside of rocks, organisms gain a range of
advantages compared to surface organisms, such
Communities; Cyanobacteria; Extremophiles; as protection from UV radiation (Cockell
Lithic habitats et al. 2003). In the Antarctic polar desert, the
stones warm the hypolithic biota and, during the
summer, mitigate freeze-thaw (Broady 1981).
Synonyms Quartz stones in the Vestfold Hills, Antarctica,
could provide temperatures 10 in excess of
Subliths ambient air temperatures (Smith et al. 2000).
For photosynthetic organisms, the limiting factor
for the thickness of rocks under which photosyn-
Definition thetic microorganisms can grow is set by the
thickness that reduces light levels to below
Hypoliths are organisms or communities of those required for ▶ photosynthesis. Investiga-
organisms that live on the underside of rocks or tions on the colonization of the underside of
at the rock-soil interface. translucent flint in the Negev desert showed that
the more translucent flint types were colonized in
a greater number of instances compared to less
Overview translucent flints (Berner and Evenari 1978). Less
than 0.01 % of incident light penetrated to below
In hot and cold deserts, the underside of rocks can 40 mm in quartz rocks in the Negev (Berner and
provide a refugium for microorganisms, both Evenari 1978), and light was reduced to 0.08 % of
photosynthetic (cyanobacteria and algae) and incident under 25 mm of quartz rock from the
non-photosynthetic (Cameron and Blank 1965; Mojave desert (Schlesinger et al. 2003). At these
Schlesinger et al. 2003). The organisms are depths colonization became limited. Broady
referred to as “hypoliths.” The community is found similarly large attenuations under Antarc-
termed “hypolithon” (following the terminology tic quartz rock (Broady 1981). The moisture
for endoliths by Golubic et al. 1981). The photo- availability in the hypolithic habitat is also an
synthetic components of hypoliths include organ- important determinant of which rocks are colo-
isms adapted to extreme rock habitats including nized and their ecological distribution in the
Chroococcidiopsis and Gloeocapsa species. environment (Warren-Rhodes et al. 2007).
Hypoliths often display well-defined “bands” of Hypoliths are also found under opaque rocks in
growth on the underside of rocks or in the case of polar deserts (Cockell and Stokes 2004). Greater
thin rocks, complete colonization of their under- than 90 % of rocks examined were colonized.
side. As photosynthetic microorganisms provide Periglacial processes can cause movements in
a source of carbon for heterotrophic microorgan- rocks, which in turn create openings around the
isms, the hypolithic colonization of translucent edges of rocks that allow the penetration of pho-
rocks by photosynthetic microorganisms can tosynthetically active radiation to the underside
indirectly benefit non-photosynthetic microor- of rocks. These processes allow for the coloni-
ganisms (Smith et al. 2000). Hypolithic zation of rocks with productivity of these
Hypoxanthine 1177
Hypolithic, Fig. 1 An
example of a community of
photosynthetic hypoliths Surface
inhabiting the underside of
opaque rocks in the
Canadian High Arctic
Colonized
underside
No colonization
(Light extinguished)
H
communities potentially as high as the above- Schlesinger WH, Pippen JS, Wallenstein MD, Hofmockel
ground productivity accounted for by plants (see KS, Klepeis DM, Mahall BE (2003) Community com-
position and photosynthesis by photoautotrophs under
Fig. 1). quartz pebbles, Southern Mohave Desert. Ecology
84:3222–3231
Smith MC, Bowman JP, Scott FJ, Line MA
See Also (2000) Sublithic bacteria associated with Antarctic
quartz stones. Antarct Sci 12:177–184
▶ Antarctica Warren-Rhodes KA, Rhodes KL, Boyle LN, Poiting SB,
▶ Cryptoendolithic Chen Y, Liu SJ, Zhuo PJ, McKay CP
(2007) Cyanobacterial ecology across environmental
▶ Extremophiles gradients and spatial scales in China’s hot and cold
▶ Photosynthesis deserts. FEMS Microbiol Ecol 61:470–482
Berner T, Evenari M (1978) The influence of temperature Hypoxanthine

and light penetration on the abundance of the
hypolithic algae in the Negev Desert of Israel. Shin Miyakawa
Oecologia 33:255–260
Broady PA (1981) The ecology of sublithic terrestrial
algae at the Vestfold Hills, Antarctica. Br Phycol
J 16:231–240
Budel B, Wessels DCJ (1991) Rock inhabiting blue-green Synonyms
algae from hot arid regions. Arch fur Hydrobiol
92:385–398
Cameron RE, Blank GB (1965) Soil studies – microflora 6-Hydroxypurine
of desert regions VIII. Distribution and abundance of
desert microflora. Space Programs Summ 4:193–202
Cockell CS, Stokes MD (2004) Widespread colonization
by polar hypoliths. Nature 431:414
Definition
Cockell CS, Rettberg P, Horneck G, Scherer K, Stokes
DM (2003) Measurements of microbial protection Hypoxanthine (C5H4N4O, molecular weight:
from ultraviolet radiation in polar terrestrial microhab- 136.11) is a naturally occurring purine base. It is
itats. Polar Biol 26:62–69
Golubic S, Friedmann I, Schneider J (1981) The
occasionally found as a constituent of nucleic
lithobiontic ecological niche, with special reference acids such as tRNA. The half-life of hypoxan-
to microorganisms. J Sed Petrol 51:0475–0478 thine to hydrolysis is 12 days at 100 C and
1178 HZE Particle
5,000 years at 0 C at pH 7. It has a UV absorption Definition

maximum at 249.5 nm (pH 7). It has been found
in the Murchison meteorite and is synthesized by HZE particles are a component of cosmic radiation
hydrolysis of ▶ adenine, in HCN polymeriza- consisting of energetic heavy nuclei (atomic num-
tions, and in discharge experiments using gas ber 3 or greater), so named for their high
mixtures such as CO–N2–H2O. The ribonucleo- (H) atomic number (Z) and high energy (E).
side of hypoxanthine is named inosine. They contribute roughly 1 % of the flux of galactic
cosmic radiation and to an even lesser fraction to
▶ solar particle events. Because of the difficulty of
See Also adequate shielding and the special nature of HZE
particle-produced lesions, these particles are
▶ Adenine considered a major hazard to living beings in
▶ HCN Polymer space, especially outside Earth’s magnetosphere,
▶ Hydrogen Cyanide that is, to human exploratory missions and
▶ Meteorite, Murchison interplanetary transfer of life by natural processes.
▶ Nucleic Acids
▶ Purine Bases
See Also
▶ Biostack
▶ Cosmic Rays in the Heliosphere
HZE Particle
▶ DNA Damage
▶ Linear Energy Transfer
Gerda Horneck
▶ Lithopanspermia
▶ Panspermia
Germany
▶ Solar Particle Events
Synonyms
Heavy charged particle; Heavy ion; Heavy

nucleus; Heavy primary
I
IAF conference on space activities. More than 1,600

technical papers on space programs and research
Michel Viso activities around the world are presented in various
CNES/DSP/SME, Vétérinaire/DVM, sessions. The IAF maintains an online archive of
Astro/Exobiology, Paris Cedex 1, France past congress papers and presentations to provide
members with a historical record of global space
activities.
Synonyms In 2010, the IAF had 205 members from
58 countries. The organizations involved include
International Astronautical Federation astronautical and other professional societies,
space agencies and international organizations,
companies from the space industry, universities
Definition and other research institutions, and nonprofit
organizations interested in space matters.
In September 1950, Alexandre Ananoff from the
“Groupement astronautique français” convened the
first International Astronautical Congress (IAC).
Eighteen representatives from the astronautical Iapetus
societies of eight countries (Argentina, Austria,
Denmark, France, Germany, Spain, Sweden, the Therese Encrenaz
UK) decided to set up an association to secure future LESIA, Observatoire de Paris - Section de
international cooperation. The International Astro- Meudon, Meudon, France
nautical Federation (IAF) was formally established
at the second IAC organized by the British
Interplanetary Society (BIS), in London, in 1951. Definition
Every year since then, the federation, together
with the International Academy of Astronautics Iapetus was discovered in 1671 by Giovanni
(IAA) and the International Institute of Space Domenico Cassini; it is the outermost midsized
Law (IISL), has organized the International Astro- icy satellite of ▶ Saturn. Its distance to Saturn is
nautical Congress, which encourages the advance- 3,560,000 km (or 59 Saturnian radii), and its
ment of knowledge about space. It is held in diameter is 1,440 km. Its density is 1.02 g/cm3,
connection with a major space exhibition. IAC is indicating a very low rock/ice ratio. Iapetus is
now recognized as the main international unique in its albedo distribution, which is very
DOI 10.1007/978-3-662-44185-5
1180 IAU
low on Iapetus’ leading side in its orbit around Overview

Saturn and ten times higher on the opposite side.
It has been proposed that the dark material could The scientific and educational activities of the
be due to the permanent accretion of matter com- IAU are organized by its nine scientific divisions
ing from the neighboring satellite ▶ Phoebe, and some 34 specialized commissions (e.g., com-
which is also very dark. Another possible expla- missions devoted to Astrobiology, History of
nation is that the low albedo of Iapetus could be a Astronomy, Stellar Evolution, The Local Uni-
result of organic deposits formed from methane verse) that cover the full spectrum of Astronomy.
ice impacted by dust particles. The long-term policy of the IAU is defined by the
triennial General Assembly and implemented by
the Executive Committee, while the day-to-day
operations are directed by the IAU officers. The
See Also
IAU Secretariat is hosted by the Institut
d’Astrophysique de Paris, France.
▶ Phoebe
The key activity of the IAU is the organization of
▶ Saturn
scientific meetings, including nine international IAU
Symposia every year, with additional symposia and
other meetings every 3 years at the General Assem-
bly. The IAU is also responsible for the definition of
IAU fundamental astronomical and physical constants,
unambiguous astronomical nomenclature, promo-
William M. Irvine tion of educational activities in astronomy, and
University of Massachusetts, Amherst, MA, USA informal discussions on the possibilities for future
international large-scale facilities. Furthermore, the
IAU serves as the internationally recognized author-
Keywords ity for assigning designations to celestial bodies and
surface features on them. The IAU is a member of
Astronomy; ICSU the International Council for Science (ICSU).
Ice
Synonyms
International Astronomical Union
Japan
Definition Institute for Advanced Study, Princeton, NJ,
USA
The International Astronomical Union (IAU) was Blue Marble Space Institute of Science,
founded in 1919 with the mission of promoting Washington, DC, USA
and safeguarding the science of astronomy in all Center for Chemical Evolution, Georgia Institute
its aspects through international cooperation. of Technology, Atlanta, GA, USA
More than 10,000 professional astronomers
(active in professional research and education)
from all over the world, at the Ph.D. level and Definition
beyond, are IAU members. The IAU is financed
by academies or equivalent institutions of its In chemistry, ice refers to water in a solid
73 member countries. crystalline phase, the common form of which
Ida 1181
encountered at normal terrestrial temperatures References and Further Reading

and pressures is known as Ice Ih. There are
16 known crystalline phases of water ice, Hobbs PV (2010) Ice physics. Oxford University Press,
New York
most of which are only known to occur at
Klinger J (1983) Extraterrestrial ice. A review. J Phys
very high pressures, pressures which may be Chem 87(21):4209–4214
encountered in the interiors of large icy Klinger J, Benest D, Dollfus A, Smoluchowski R (eds)
planets or moons, although, one other phase, (1985) Ices in the solar system. Springer, New York
Ice XI has been observed in some Antarctic
glaciers. Ice Ih is the only known nonmetallic
substance which expands as it freezes, being
some 9 % less dense than liquid water at Ice Line
273 K. This is due to hydrogen bonding in the
crystal lattice which forces the molecules into ▶ Snow Line
positions farther apart than normally occurs in
the liquid phase. This change in volume contrib-
utes to water’s activity in the mechanical
weathering of minerals and also causes ice to Ice-Albedo Instability
I
float when frozen.
In planetary science, and more generally in ▶ Snowball Earth
astronomy, an ice refers to a volatile (a species
with a low boiling point, such as N2, water,
CO2, NH3, H2, CH4, and SO2) with a
melting point above 100 K. The solid 243 IDA
phases of these ices can be glasses (which are
technically extremely viscous liquid phases) or ▶ Ida
crystalline solids, depending on the temperature,
pressure, and rate at which the ice forms.
Ices observed or expected to be present on the
mantles of interstellar dust grains include, in Ida
addition to water, CO, CO2, CH3OH, CH4, NH3,
O2, and N2. Ices reported on the surfaces of sat- Ivanka Pelivan
ellites in the outer solar system and of Kuiper Institute of Planetary Research, German
Belt Objects include H2O, CH4, N2, CO, Aerospace Center, Berlin, Germany
CH3OH, and perhaps other volatile organics
such as ethane.
Synonyms
243 IDA
See Also
▶ Comet Definition
▶ Enceladus
▶ Europa Ida is a main-belt asteroid of the Koronis family,
▶ Hydrogen Bond originating from the breakup of a larger parent
▶ Interstellar Ices body. It orbits the sun between the orbits of Mars
▶ Kuiper Belt and Jupiter at an average distance of 2.86 AU
▶ Volatile with one orbit taking about 4.84 years. Ida has a
▶ Water rotation period of 4.63 h. Elongated in shape, its
1182 IDP
average diameter is about 32 km. It is accompa-

nied by its moon Dactyl which has an average IEP
diameter of 1.2–1.6 km. Dactyl’s orbit provided
constraints for Ida’s density to 2.6 0.5 g/cm3. ▶ Isoelectric Point
Ida’s mass is estimated between 3.65 and
4.99 1016 kg.
Ida has been classified as spectral S-type aster-
oid. The irregular surface of the asteroid is Igneous Rock
heavily cratered and covered by regolith com-
posed of silicate minerals. Ida exhibits space Nicholas Arndt
weathering where older parts of the surface turn ISTerre, Université Grenoble Alpes, France
reddish. Based on cratering records, Ida’s surface
seems more than 1 billion years old, whereas the
Ida-Dactyl system is estimated to be less than Synonyms
100 million years of age considering the colli-
sional life expectancy of Dactyl. Magmatic rock
History Definition
Ida was discovered in 1884 by the Austrian An igneous rock forms by solidification of
astronomer Johann Palisa in Vienna. It was vis- magma. There are two main types: volcanic
ited and imaged in 1993 by the Galileo space- rocks that crystallize or solidify from erupted
craft. Ida’s moon Dactyl, the first confirmed lava or fragmental (pyroclastic) material that
satellite of an asteroid, was discovered in Febru- reached the surface and contacted the atmo- or
ary 1994 by Ann Harch from Galileo’s hydrosphere and plutonic or intrusive rocks that
imaging team. crystallize from magma that did not reach the
surface. These two types are distinguished by
their grain size: intrusive rocks are coarse-
See Also grained and typically show an interlocking fabric
because they crystallize slowly in a thermally
▶ Asteroid insulated setting, whereas volcanic rocks are
▶ Asteroid Belt, Main fine-grained or even glassy because they cool
▶ Galileo Galilei rapidly following eruption at the Earth’s surface;
▶ Regolith, Planetary crystals typically float in a fine-grained matrix.
Common minerals in igneous rocks are olivine,
pyroxene, amphibole, mica, feldspar, and quartz.
Common intrusive rock types include peridotite,
▶ gabbro, granodiorite, and ▶ granite; their vol-
IDP
canic equivalents are komatiite, ▶ basalt, andes-
ite, and rhyolite, respectively.
See Also
IEC ▶ Basalt
▶ Cryovolcanism
▶ Ion-Exchange Chromatography ▶ Gabbro
Ikhwan al-Safa 1183
▶ Granite sometimes called books: “mathematical sci-

▶ Hydrothermal Environments ences” (14 epistles), “corporeal and natural sci-
▶ Mafic and Felsic ences” (15 epistles), “sciences of the soul and
▶ Mantle intellect” (ten epistles), “Divine and legal sci-
▶ Mid-ocean Ridges ences” (11 or 12 epistles). Ikhwān al-Safā
˙
▶ MORB addressed astronomical and cosmological con-
▶ Plate Tectonics siderations in the second book, especially in its
▶ Volcano second epistle (16 overall). Ikhwān al-Safā
˙
endorsed the concept of the celestial animal,
which Ptolemy presented in his Planetary
Hypotheses, and which had important Stoic
roots. Similar to an animal or a human, the uni-
Ikhwan al-Safa
verse with its different orbs is animated by the
universal soul, which leaves it at the end of times,
1 much like the departure of a human or animal
Harvard Divinity School, Cambridge, MA, USA
2 soul signals the end of life for that given being.
Harvard University, Cambridge, MA, USA
Their discussion of the universal soul, and the end
I
of the universe at the end of times, departed from
Aristotelian views on the infinity of the universe
Keywords
which were common among philosophers at the
time, and share the convictions of theologians
Islam; Islamic astronomy; Isma’ili; Fatimid;
(mutakallimī n).
Pythagorean; Celestial animal; Astrology
Although Ikhwān al-Safā’s writings betray
˙
strong Pythagorean and Neo-Platonic influences,
their cosmological and astronomical discussions
Synonyms largely accepted Aristotelian and Ptolemiac
models that were dominant in the Greco-Islamic
Brethren of Purity context of the time. They accepted the universal
circular motion of planets as caused by the move-
ment of the outer orb, which is moved by the
Overview Prime Mover. Motion was transferred down-
wards from outer to inner orbs, making lower
The Brethren of Purity (Ikhwan al-Safa’) were a planets slower than the upper planets. They
˙
group of anonymous authors who composed a unequivocally refused any notion of contradic-
collection of 50–52 epistles (the last two may tory or retrograde motion, insisting that all
have been later additions by the same or other planets moved in the same direction in perfect
authors), which became well-known from the late circles. They appeared to have objected to
ninth century. Contemporary Shiite Ismaʿīlī Ptolemy’s eccentrics, although they fully
scholars and authors claimed the epistles to be endorsed epicycles as perfect explanations of
composed by Ismaʿīlī missionaries—a claim that the planets’ retrograde motion and of apparent
went uncontested by their contemporaries. This change in their size and brightness. Ikhwān
Ismaʿīlī attribution combined with internal evi- al-Safā accepted the Aristotelian notion of a dif-
˙
dence suggest that the epistles were either fully or ferent nature for celestial beings. However, they
largely completed before the establishment of the explained that this nature was limited only to
Ismāʿīlī Caliphate in 909. their being incorruptible, and that they exhibited
As a collection, the Epistles attempted to many of the other qualities of bodies in the sub-
encompass all branches of knowledge. The lunar sphere. These similarities appeared to pro-
52 Epistles were grouped into four sections, vide enough justification for celestial influences
1184 IKI
on beings in the sublunar sphere, thus legitimiz- Definition

ing astrology. They rejected the existence of
empty space whether inside or outside the IKI, the Russian Space Research Institute
universe. (Russian: Инcтитут кocмичecкиx
While the Epistles represented an encyclope- иccлeдoвaний Poccийcкoй Aкaдeмии Haук), is
dic collection that addressed all essential an institute of the Russian Academy of Sciences
branches of knowledge at the time, they had a that was founded in 1965 as the Space Research
central goal of self-improvement that held Institute of the USSR Academy of Sciences and
through the entire collection. They also renamed in 1992. It is the leading Russian orga-
maintained a clear understanding of prophecy nization for space research and has frequently
and the Imamate and their role as conduit of collaborated on missions with ▶ NASA, ▶ ESA,
universal divine knowledge, the difference and other national space agencies. IKI is located
between the Select and the Commoners in rela- in Moscow and has a staff of some 290 scientists
tion to knowledge, and the role of knowledge as in the fields of astrophysics, planetary sciences,
the central component of faith and pious living. solar-terrestrial physics, cosmic plasma physics,
and geophysics. Among its successes relevant to
astrobiology was the ▶ Vega spacecraft to
▶ Comet Halley.
De Callataÿ G (2005) Ikhwan al-Safa’: a brotherhood of

idealists on the fringe of orthodox Islam. Oneworld
Publications, Oxford See Also
Nasr SH (1964) An introduction to Islamic cosmological
doctrines: conceptions of nature and methods used for ▶ Vega 1 and 2 Spacecraft
its study by the Ihwan Al-Safa, Al-Biruni, and Ibn
Sina. Harvard University Press, Cambridge, MA
Netton IR (2002) Muslim Neoplatonists: an introduction
to the thought of the brethren of purity (Ikhwān
al-Safāʾ). Routledge Curzon, New York
˙
The series “Epistles of the Brethren of Purity,” edited by Imaging
Nader El-Bizri and published jointly by Oxford
University Press and the Institute of Ismaili Studies
since 2008, will be the first critical edition and
Daniel Rouan
annotated English translation of the entire corpus of LESIA, Observatoire Paris-Site de Meudon,
the Rasāʾil, gradually replacing the three previous Meudon, France
editions of the text (Bombay 1887–9, Cairo 1928,
Beirut 1957)
Keywords
Adaptive optics; CCD; Camera; Direct detection;

Telescope
IKI
Michel Viso
CNES/DSP/SME, Vétérinaire/DVM, Definition
Imaging is the set of techniques used for
obtaining images of one or several astronomical
Synonyms bodies or fields using a two-dimensional detector,
computerized techniques, and smart optical tech-
Russian Space Research Institute niques such as ▶ adaptive optics.
Imaging 1185
Overview
In astronomy, imaging is in most cases the first

observational approach to discovering a new
object, the image being required to clarify struc-
ture and morphology, before using the more
physical tool of ▶ spectroscopy. Shape, size,
decomposition into elements, arrangement of
subparts, measurement of displacements, and dis-
crimination of photometric variations belong to
the set of information provided by the image.
From the planets in the solar system to the iden-
tification of gigantic filamentary structures or
sheets in the universe, the image has played a
major role in astronomy. Improving the quality Imaging, Fig. 1 The Megacam camera developed by
of the image has been and remains today more CEA-France and installed at the prime focus of the
than ever a powerful drive in instrumental Canada-France-Hawaii Telescope, behind a wide field
corrector. This huge camera includes 36 CCDs of I
research: two-dimensional detectors, high angu- 2,048 4,612 pixels each, arranged side by side, leading
lar resolution, ▶ adaptive optics, high dynamical to a total of 340 Mpixels
range, and post-processing techniques are fields
of intense activity. One notes that the direct
detection of exoplanets is certainly among the suppress artifacts. Thanks to the ▶ adaptive
most demanding topics in terms of angular reso- optics technology that can negate to a large extent
lution and high contrast. the atmospheric blurring, image quality can
Detectors: In the last 30 years, the photo- approach the theoretical resolution capability of
graphic technique has been largely replaced the telescope, that is, angular details as small as
by digital sensors such as CCDs and now y = l/D, where D is the diameter of the telescope
CMOS chips (complementary metal-oxide- and l the wavelength. For l = 2 mm and a
semiconductor, a technology used for D = 8 m, this corresponds to a length of 10 km
constructing integrated circuits). Giant cameras on Venus’ surface or the orbit of Mercury seen
associating several tens of CCD chips can feature around a star at 300 light-years. In space, almost
up to 500 Mpixels and cover extra large fields on perfect conditions also provide excellent angular
the sky, as demonstrated, for instance, by resolution, despite a more moderate telescope
Megacam, a camera installed at the Canada- size, as illustrated by the stunning images of the
France-Hawaii Telescope (Fig. 1). CCDs provide Hubble Space Telescope. The future James Webb
numerical images, which can easily be processed Space Telescope with its 6.5 m diameter tele-
by computers. They also offer a very good detec- scope will soon provide similar exquisite image
tion efficiency in a large domain of wavelength. quality but in the infrared domain.
Beyond l = 1 mm, infrared arrays based on the As regards high contrast imaging, the recent
CMOS technique have about the same advan- pressure put by the search for exoplanets was the
tages. Recently large arrays of infrared bolome- cause of an explosion of ideas and new concepts
ters have been produced, allowing imaging in to mitigate the problem of the huge difference in
even the submillimeter domain. brightness between a star and the planets orbiting
A certain degree of computer correction for it. New types of coronagraphs to hide the star-
atmospheric and instrumental effects allows light, of wavefront control to reduce speckles,
sharpening up the image and improving the con- and of pupil shaping to suppress the wings of
trast. Multiple digital images can also be com- the point spread function produced by diffraction
bined to further enhance the signal-to-noise and have been proposed and tested. Also, differential
1186 2, 4-Imidazolelidinedione
Léna P, Rouan D, Lebrun F, Mignard F, Pelat D (2011)

Observational astrophysics: astronomy and astrophys-
ics library. Springer, Berlin, XV + 719 p. ISBN 978-3-
642-21815-6
Starck J-L, Murtagh F (eds) (2006) Astronomical image
and data analysis. Astronomy and astrophysics library.
Springer, New York
2, 4-Imidazolelidinedione
▶ Hydantoin
Imaging, Fig. 2 High-contrast imaging. Thanks to the

combination of dedicated optical devices (here a four- Imidogen
quadrant phase mask coronagraph) and imaging tech-
niques (here differential rotation), it is possible to reach William M. Irvine
an extremely high contrast ratio at the laboratory: a fake
planet 0.15 billion times fainter than an artificial star University of Massachusetts, Amherst, MA, USA
(behind the central occulted disk) is perfectly seen on
this image obtained at LESIA, Observatoire de Paris
Synonyms
techniques based on chromatic differences, or on NH; Nitrogen hydride
image rotation, or on polarization properties have
been refined and greatly improve the detection
capability. Today in the laboratory, a contrast of
109 is reached at an angular distance of a few Definition
l/D only (Fig. 2). The next decade will see those
techniques implemented in ground-based or This diatomic radical, containing nitrogen and
space instruments. hydrogen, has been detected in both the diffuse
▶ interstellar medium, where it is seen in absorp-
tion against the light of background stars, and in
See Also denser gas toward the center of our ▶ Milky Way
galaxy. In the latter case the relative abundances
▶ Adaptive Optics of NH3, NH2, and NH have been interpreted in
▶ CCD terms of low-velocity shock activity (Goicoechea
▶ Coronagraphy et al. 2004).
▶ Telescope
Aime C (2007) Optical techniques for direct imaging of NH was first detected at near ultraviolet wave-
exoplanets. CR Phys 8:298 lengths in 1991 by D. M. Meyer and K. C. Roth.
Howell SB (2006) Handbook of CCD astronomy: Cam-
bridge observing handbooks for research astronomers,
The subsequent observations of the Galactic cen-
vol 5, 2nd edn. Cambridge University Press, Cam- ter were made with the Infrared Space Observa-
bridge. ISBN 0-52185-215-3 tory (ISO). It has been recently observed using
Impact Degassing 1187
the Herschel satellite towards other ▶ molecular Caloris Basin on ▶ Mercury, the Orientale Basin
clouds (Persson et al. 2010, 2012). on the Moon, and the Hellas Basin on ▶ Mars.
See Also
See Also
▶ Milky Way
▶ Crater, Impact
▶ Facula, Faculae
▶ Mars
References and Further Reading ▶ Mercury
▶ Moon, The
Goicoechea JR, Rodrı́guez-Fernández NJ, Cernicharo
J (2004) The far-infrared spectrum of the sagittarius
B2 region: extended molecular absorption, photodis-
sociation, and photoionization. Astrophys
J 600:214–233
Impact Degassing
Persson CM, Black JH, Cernicharo J et al (2010) Nitrogen I
hydrides in interstellar gas: Herschel/HIFI observa-
tions towards G10.6-0.4 (W31C). Astron Astrophys Reika Yokochi
521:L45 Department of Geophysical Sciences,
Persson CM, De Luca M, Mookerjea B et al (2012) Nitro-
gen hydrides in interstellar gas. II. Analysis of Her- The University of Chicago, Chicago, IL, USA
schel/HIFI observations towards W49N and G10.6-0.4 Department of Earth and Environmental
(W31C). Astron Astrophys 543:145 Sciences, University of Illinois at Chicago,
Chicago, IL, USA
Impact Basin Keywords
Roland J. Wagner Planetary accretion; Meteor impacts; Formation

German Aerospace Center (DLR), Institute of of planetary atmospheres; Volatile inventory of
Planetary Research, Berlin, Germany planets
Definition Synonyms
An impact basin is a large complex impact crater. Shock devolatilization

The threshold diameter to distinguish
between craters and basins is approximately
150–200 km. In general, basins are characterized Definition
by two or more concentric rings, which are ridges
or scarps facing toward the basin. One of these Impact degassing refers to the release of gas
rings is the main rim that borders the cavity from caused by a high-velocity collision of volatile-
which material was excavated and ejected during bearing solid objects, particularly during colli-
the impact. Most basins are heavily degraded or sions between planetary objects. During an
have been covered by younger material. All event or collision, an intense mechanical shock
known basins are old impact features created wave is transmitted, causing changes in specific
during the first 800 million years of the planet volumes along the material-specific Hugoniot
or satellite on which they are found. Examples are curve (see ▶ Spallation Zone). A rarefaction
1188 Impact Degassing
wave subsequently propagates to reduce the directing the surface evolution of planets and the
shock pressure, during which the kinetic energy chemical compositions of planetary atmospheres.
of the collision is released as heat and as high Recent shock-loading experiments and theoretical
particle velocity of the ejecta. The change in studies involve detailed investigation of chemical
thermodynamic conditions causes phase transi- changes, for example, on hydration of olivine by
tions of the material in the colliding objects, shock-released water (Furukawa et al. 2011), on
either in the solid state or by melting and vapor- the survival or formation of amino acids that have
ization, which releases structurally bonded vola- direct implications for life-forming molecules
tiles as well as interstitially trapped gases. (Pierazzo and Chyba 1999; Sugita and Schultz
2009), or destruction of complex molecules to
form chemically simpler constituents of atmo-
Overview spheres (Fukuzaki et al. 2010).
The moon-forming impact caused large-scale
The discovery of impact craters on the Moon melting of the Earth, resulting in a large-scale
during the Apollo missions first inspired the stud- degassing from the silicate Earth to the atmo-
ies of collisional shock on geological materials. sphere. Whether this event resulted in significant
Theoretical studies started by investigating the loss of atmosphere to space and/or chemical and
thermodynamic stabilities of mineral matrices in isotopic fractionation of the terrestrial atmo-
colliding geological objects that are free of struc- sphere remains controversial.
tural volatiles (Ahrens and O’Keefe 1972; Luo
and Ahrens 2004).
In the context of planetary evolution and the See Also
formation of planetary atmospheres that may
have fostered emergence of life, the main focus ▶ Crater, Impact
of investigation concerns the release of structur- ▶ Earth’s Atmosphere, Origin and Evolution of
ally bound major volatiles from hydrous min- ▶ Oceans, Origin of
erals, carbonate, as well as ice. The shock- ▶ Water, Delivery to Earth
loading experimental studies demonstrated that
structurally bound water in hydrous minerals References and Further Reading
(Boslough et al. 1980; Lange et al. 1985) could
be lost during hypervelocity impacts, whereas Ahrens TJ, O’Keeffe JD (1972) Shock melting and vapor-
theoretical and experimental studies suggest that ization of lunar rocks and minerals. In: Conference on
minerals containing volatile elements as major lunar geophysics, p 214–249
Boslough MB, Weldon RJ, Ahrens TJ (1980) Impact-
constituents (i.e., carbonate, sulfate, and H2O induced water loss from serpentine, nontronite and
ice) initiate vaporization at relatively kernite. In: Lunar and planetary science conference
low-impact velocity (Shen et al. 2003; Stewart XI, Houston, p 2145–2158
and Ahrens 2005; Ohno et al. 2008). Tyburczy Fukuzaki S, Sekine Y, Genda H, Sugita S, Kadono T,
Matsui T (2010) Impact-induced N2 production from
et al. (1986) experimentally investigated shock- ammonium sulfate: implications for the origin and
induced volatile loss from the Murchison CM2 evolution of N2 in Titan’s atmosphere. Icarus
chondrite and observed that complete devolati- 209(2):715–722. doi:10.1016/j.icarus.2010.04.015
lization occurs at a pressure of about 30 GPa, Furukawa Y, Sekine T, Kakegawa T, Nakazawa H (2011)
Impact-induced phyllosilicate formation from olivine
corresponding to ~27 % of Earth’s present mass. and water. Geochim Cosmochim Acta
This validates that the conclusions based on indi- 75(21):6461–6472. doi:10.1016/j.gca.2011.08.029
vidual minerals above generally apply when they Lange MA, Lambert P, Ahrens TJ (1985) Shock effects on
are embedded with other minerals in the primitive hydrous minerals and implications for carbonaceous
meteorites. Geochim Cosmochim Acta 49:1715–1726
meteorite, a potential precursor of planets. It is Luo S-N, Ahrens TJ (2004) Shock-induced superheating
therefore likely that impact degassing from plane- and melting curves of geophysically important min-
tary precursor materials played a significant role in erals. Phys Earth Planet Inter 143–144:369–386
Impact Parameter 1189
Pierazzo E, Chyba CF (1999) Amino acid survival in large generated during the impact. The largest craters
cometary impacts. Meteorit Planet Sci 34(6):909–918. such as Sudbury (Ontario, Canada), Chicxulub
doi:10.1111/j.1945-5100.1999.tb01409.x
Stewart ST (2005) Shock properties of H2O ice. J Geophys (Mexico), Popigai (Siberia), or Manicouagan
Res 110(E3):E03005. doi:10.1029/2004JE002305 (Québec, Canada) contain a very large volume of
Sugita S, Schultz PH (2009) Efficient cyanide formation impact melt. In the case of Sudbury, it also hosts
due to impacts of carbonaceous bodies on a planet with important ore deposits of nickel.
a nitrogen-rich atmosphere. Geophys Res Lett 36(20):
L20204. doi:10.1029/2009GL040252
Tyburczy JA, Frisch B, Ahrens TJ (1986) Shock-induced
volatile loss from a carbonaceous chondrite: implications
for planetary accretion. Earth Planet Sci Lett 80:201–207 See Also
▶ Chicxulub Crater
▶ Crater, Impact
▶ Gunflint Formation
Impact Melt Rock
▶ Impactite
Philippe Claeys
Earth System Science, Vrije Universiteit Brussel,
Brussels, Belgium I
Impact Parameter
Definition Daniel Rouan
An impact melt rock is a type of rock typically Meudon, France
found in impact structures. It is an ▶ impactite
consisting of a mixture of solid and melt fragments
all derived from the target rock, floating in a Definition
glassy, microcrystalline, or recrystallized matrix.
The solid fragments may contain shocked min- The impact parameter is the projected distance
erals. An impact melt rock results from the cooling between the axis carrying the initial vector veloc-
of the melt pool that formed within a crater ity of a projectile and the center of the target that
because of the high temperature and pressure the projectile is approaching (see Fig. 1).
Impact Parameter,
Fig. 1 Definition of the
impact parameter (dark
arrow) of a projectile
directed toward a target
1190 Impact, Hit and Run
Impact, Hit and Run Impactite
Sean N. Raymond Philippe Claeys

Laboratoire d’Astrophysique de Bordeaux, Earth System Science, Vrije Universiteit Brussel,
CNRS, Universite de Bordeaux, France Brussels, Belgium
Definition
Definition
During terrestrial planet formation, the so-called
hit-and-run impacts are giant impacts that do not Impactite is a term used to describe rocks formed
result in a net increase in the mass of at least one during a meteorite impact on a planetary body.
of the planetary embryos involved in the colli- The term covers the lithologies such as ▶ breccia,
sion. Hit-and-run impacts generally imply high ▶ suevite, or ▶ impact melt rock found within or
relative velocities between bodies and/or oblique at close proximity of an impact structure.
impact angles. In such cases, the larger body does Monomictic breccias contain fragments of a
not accrete the smaller body, but the larger object single rock type; polymictic breccias contain
could actually be eroded and sometimes severely several rock types. Each is composed of
traumatized by the encounter. A significant chaotic assemblages of solid and shocked frag-
amount of debris may be created in hit-and-run ments, or previously molten fragments, of the
impacts. target lithologies. Suevite is a breccia containing
melt and solid particles surrounded by a fine
clastic matrix, first defined in the Ries crater
(Germany). Distal impactite, composed of fine
See Also ejected material (spherules, shocked minerals),
may be widespread in the case of large impacts
▶ Giant Impact such as the one that formed the ▶ Chicxulub
▶ Late-Stage Accretion crater.
See Also
Impact, Probability ▶ Breccia

▶ Chicxulub Crater
Catharine A. Conley ▶ Crater, Impact
NASA Headquarters, Washington, DC, USA ▶ Impact Melt Rock
▶ KT Boundary
▶ Suevite
Definition
For purposes of ▶ planetary protection, the prob-

ability of impact is the probability, determined by
a sequential evaluation of possible spacecraft
failure modes, that a spacecraft might impact a In Vitro Evolution
planetary body under nominal and off-nominal
conditions. ▶ Evolution, In Vitro
Inelastic Photon Scattering 1191
is especially important in the ▶ radial velocity

Inactivation technique used for detecting exoplanets, since
the derived planetary mass is undetermined by a
Catharine A. Conley factor 1/sin (i): if the orbital plane is close to the
NASA Headquarters, Washington, DC, USA plane of the sky, the actual mass can be much
larger than the raw value.
Definition
In ▶ planetary protection, the term “inactivation” See Also

expresses the result of a process that
renders microorganisms incapable of reentering ▶ Orbit
a proliferative state. The parameters (time, ▶ Radial Velocity
temperature, concentration, dose, etc.) of a pro-
cess which allows inactivation of 90 % of the
population of a given microorganism define the
▶ D-value.
Indeterminacy I
▶ Chance and Randomness

See Also
▶ Disinfection
▶ D-Value Indian Space Research Association
▶ Pasteurization
▶ Sterilization ▶ ISRO
Indian Space Research Organization

Inclination (Astronomy)
▶ ISRO
Daniel Rouan
Meudon, France
Indigenous
Definition
▶ Endogenicity
The inclination of an object in the solar system is
the angle between the orbital plane of the object
(planet, comet, or asteroid) and the plane of the
▶ ecliptic.
In binary systems and especially in star/planet Inelastic Photon Scattering
systems, it corresponds to the angle i between the
orbital plane and the plane of the sky. This angle ▶ Raman Scattering
1192 Infrared Astronomical Satellite
UK. Launched in January 1983, it produced infra-

Infrared Astronomical Satellite red data for a period of ten months and covered
more than 96 % of the sky.
Molecular Astrophysics, Iorrejón de Ardoz,
Madrid, Spain See Also
▶ Herschel Mission
▶ Infrared Astronomy
Synonyms ▶ ISO
▶ Spitzer Space Telescope
IRAS

Definition http://irsa.ipac.caltech.edu/IRASdocs/iras.html
http://www.sron.rug.nl/irasserver/irasserverman.html
The Infrared Astronomical Satellite or IRAS was the
first observatory to perform an all-sky survey at
infrared wavelengths. It was equipped with a tele-
scope of 60 cm diameter and with detectors operating Infrared Astronomy
at wavelengths of 12, 25, 60, and 100 mm with
angular resolution of 30 arc sec and 2 arc min at Daniel Rouan
12 and 100 mm, respectively. Launched January LESIA, Observatoire Paris-Site de Meudon,
25, 1983, it was operating up to November Meudon, France
21, 1983. During its primary mission, which was to
obtain a complete map of the sky, it discovered
350,000 point-like sources, increasing the number Synonyms
of cataloged astronomical sources by about 70 %.
IRAS discoveries included a disk of dust grains Far infrared (far IR); Near infrared (near IR)
around the star Vega, six new comets, and very
strong infrared emission from interacting galaxies,
as well as wisps of warm dust called infrared cirrus Definition
which are found in almost every direction of space.
IRAS also revealed for the first time the core of our Infrared astronomy is the branch of observational
galaxy, the Milky Way. In addition to the infrared sky astronomy which studies celestial objects
maps, IRAS was also equipped with a low-resolution through their infrared radiation, that is, at wave-
spectrometer that was used to observe thousands of lengths longer than 0.75 mm and shorter than
objects in the mid-infrared. The outstanding scientific 400 mm. The domain is not strictly defined, and
results obtained by IRAS opened a new view of the some astronomers consider that it begins at 1 mm
universe and motivated new infrared space facilities where ▶ CCDs, the most universally used detec-
such as ▶ ISO, ▶ Spitzer, and ▶ Herschel. tors in the visible domain, are no longer sensitive.
Infrared astronomy is considered as part of opti-
cal astronomy because technologies and methods
History (incoherent detection, classical optics, solid-state
digital detectors) are globally the same as in the
The project was initiated in 1975 as a joint pro- visible domain. One peculiarity of infrared
gram of the USA, the Netherlands, and the astronomy is the strong ▶ background emission
Infrared Excess 1193
1.0
Transmission
0.8
0.6 14 km (Aircraft)
0.4
0.2 4 km (Mauna-Kea)
0.0
0.2 1 10 100 1,000
Wavelength (μm)
Infrared Astronomy, Fig. 1 Transmission of the atmo- particularly with telescopes on the top of high mountains
sphere in the infrared range. The Earth’s atmosphere (Mauna Kea is a summit at 4,200 m on the island of
absorbs the infrared light in the largest part of the wave- Hawaii, the best infrared site today). At higher altitudes
length domain: only in a few windows is the transmission (aircraft, balloon), the conditions are improving and
good enough to allow observations from the ground, broader windows are open
of the environment (atmosphere, telescope) that

requires dedicated differential techniques. Infrared Excess
Another one is the absorption by the atmosphere
(see Fig. 1) in a large fraction of the wavelength Steven W. Stahler I
domain: this explains why balloons, airplanes, Department of Astronomy, University of
and satellites are often used to carry instruments. California, Berkeley, CA, USA
▶ IRAS, ISO, and ▶ Spitzer are the space instru-
ments that brought the most spectacular results in
the field. Main targets in infrared astronomy are Keywords
cold objects such as cool stars (▶ brown dwarves,
M dwarves), ▶ interstellar dust, planets of the Interstellar dust
solar system, and now exoplanets.
The infrared domain is of special importance Definition
to study planets, since it’s where ▶ thermal emis-
sion peaks and molecules exhibit many spectral Many ▶ pre-main sequence stars emit substantial
features corresponding to transitions between amounts of energy at infrared and longer wave-
ro-vibrational levels of energy. ▶ Infrared spec- lengths. To the degree that the flux at each wave-
troscopy is the major tool to study the planetary length exceeds the corresponding blackbody value
atmosphere and surface. (corresponding to the temperature of the stellar
Note that the infrared range is generally photosphere), the star is said to have an infrared
subdivided into three domains: near infrared (near excess. At least half of T Tauri stars have infrared
iR), l = 1–5 mm; mid-infrared (mid-IR), excesses. Much of this emission arises from heated
l = 5–25 mm; and far infrared (far IR), l > 25 mm. dust grains embedded in circumstellar disks. Other
T Tauri stars have no infrared excess. Presumably,
these stars have no disks, although they are of
See Also similar age. All stars must lose their disks by the
time they reach the ▶ main sequence, whose stars
▶ Blackbody do not have an infrared excess.
▶ Electromagnetic Spectrum
▶ Infrared Astronomical Satellite
▶ Infrared Space Observatory Overview
▶ Infrared Spectroscopy
▶ Spitzer Space Telescope Stars radiate over a broad range of wavelengths.
▶ Thermal Emission A plot of radiation intensity as a function of
1194 Infrared Space Observatory
wavelength, when covering a substantial range in true protostars, objects deriving their luminosity
the latter, is known as a spectral energy distribu- from the infall of cloud material onto the stellar
tion. In such a plot, each intensity is measured surface.
using a relatively broad filter, which admits radi-
ation over a wavelength range of roughly 1,000 Å
(100 nm). See Also
A main sequence star radiates like a black-
body. That is, its spectral energy distribution ▶ Interstellar Dust
resembles, at least approximately, the Planck ▶ Pre-Main-Sequence Star
curve corresponding to the star’s surface temper- ▶ Protostars
ature. The spectral energy distributions of young ▶ T Tauri Star
stars, however, depart sharply from the Planck
curve. These objects are often surrounded by
considerable amounts of interstellar dust, which
absorbs and reddens the radiation in a Beckwith SVW (1999) In: Lada CJ, Kylafis ND (eds) The
wavelength-dependent manner. The raw intensi- origin of stars and planetary systems. Reidel, Dor-
ties must therefore be corrected, a procedure drecht, p 579
known as dereddening. Bertout C (1989). Ann Rev Astron Astrophys 27:351
Even after dereddening, many T Tauri stars
exhibit substantial flux above a blackbody at
infrared and longer wavelengths. In these
so-called classical T Tauri stars, the total excess Infrared Space Observatory
flux emitted in the infrared can be a substantial
fraction of the total stellar luminosity. Other Martin F. Kessler
T Tauri stars, the so-called weak-lined class, European Space Agency (ESA), European Space
have no such excess. Astronomy Centre (ESAC), Madrid, Spain
T Tauri stars are pre-main sequence objects of
roughly solar mass and below. That is, they are
contracting slowly and generating energy Keywords
through that contraction, but not yet capable of
fusing hydrogen into helium. Detailed analysis of ISO; Infrared astronomy; Organic molecules in
the infrared excess emission from classical space; Space astronomy; Satellite; Water
T Tauri stars shows that it arises from radiation
by heated dust grains. These grains must be rela-
tively close to the star. Most are embedded in Synonyms
circumstellar disks, which have been imaged
directly in some cases. It is surprising that ISO
weak-lined T Tauri stars, which have similar
ages as the classical ones, apparently lack these
disks. Definition
Pre-main sequence objects of larger mass,
from about 2 to 10 solar masses, are known as The Infrared Space Observatory (ISO), a project
Herbig Ae and Be stars. Many of these also of the European Space Agency (ESA) with the
exhibit large infrared excesses. There are also participation of ▶ NASA and ISAS (now
many stars, presumably of lower mass, which ▶ JAXA), was the first cryogenic spaceborne
are so deeply embedded in dust that all their infrared observatory for astronomy. It operated
emission is at infrared and longer wavelengths. from 1995 to 1998 and made detailed imaging
Some of these latter sources are believed to be and photometric and spectroscopic observations
Infrared Space Observatory 1195
Infrared Space
Observatory,
IR path
Fig. 1 Schematic of ISO
Sunshield with Payload module
solar cells (cryostat)
Startrackers
Service module
(for electrical power,
attitude control and
Superfluid
telecommunication)
helium tank
Telescope with
scientific instruments
Interface with + star sensor
ariane
I
at wavelengths from 2 to 200 mm of all kinds of 2.5–240 mm with spatial resolutions ranging
astronomical objects from ▶ planets and from 1.5 to 90 arc s, depending on wavelength.
▶ comets in our own Solar System right out to The Service Module provided the traditional
distant extragalactic sources. Its contributions to spacecraft services including three-axis attitude
astrobiology include the detection in space of stabilization at the arc second level (Kessler
▶ water, ▶ complex organic molecules, and a et al. 1996).
rich variety of ices (Fig. 1). The mission was a technical, operational, and
scientific success with most satellite systems
operating far better than specifications and with
Overview its scientific results impacting practically all
fields of astronomy.
The Infrared Space Observatory (ISO), selected During its 29-month operational lifetime, ISO
by ESA in 1983, made a detailed exploration of made over 30,000 individual observations. These
the infrared universe. Industrial design and devel- data are publically available at http://iso.esac.esa.
opment started in 1986 with Aérospatiale (now int/ida/. ISO’s scientific legacy is summarized in
part of Thales Alenia Space) as prime contractor. various overviews and compilations (e.g., First
An Ariane 44P vehicle launched ISO in Novem- ISO Results 1996; Kessler and Cox 1999;
ber 1995, and it operated until May 1998 in an Cesarsky and Salama 2005). The mission made
elliptical 24-h orbit with perigee/apogee of 1,000/ several significant contributions to astrobiology.
70,000 km. ISO’s Payload Module was essen- ISO is found water everywhere in the
tially a large cryostat, containing 2,300 l of super- universe – in comets, ▶ Mars, the four giant
fluid liquid helium, cooling four scientific planets and ▶ Titan, the interstellar medium
instruments and a telescope of Ritchey-Chrétien (toward protostars and molecular clouds),
design with a 60 cm diameter primary mirror to evolved stars (both oxygen and carbon rich),
temperatures of 2–4 K. The two ▶ spectrometers and ▶ galaxies (e.g., Cernicharo and Crovisier
(Short-Wavelength Spectrometer [SWS], Long- 2005; Encrenaz 2008). ISO’s wide and
Wavelength Spectrometer [LWS]), a camera uninterrupted wavelength coverage, free of tellu-
(ISOCAM), and an imaging photopolarimeter ric contamination, permitted it to make a detailed
(ISOPHOT) jointly covered wavelengths of inventory of interstellar ices (including not only
1196 Infrared Spectroscopy
the ubiquitous CO2 but also 13CO2, CO, H2O, Definition

H2CO, CH4, CH3OH, CH4, OCS, etc.) and gas
phase organic molecules (including H2O, CH4, Infrared spectroscopy is spectroscopy using
C2H2, CH3, HCN, CO2, OH, etc.) as well as infrared light (0.7–1,000 mm;
addressing the rich variety of ▶ polycyclic aro- 14,000–10 cm1). Most molecular bonds are
matic hydrocarbon features and making an inven- characterized by the absorption of specific fre-
tory of the reservoirs of the major elements (C, O, quencies of infrared radiation. The 0.7–2.5 mm
N, etc.) (e.g., Ehrenfreund and Charnley 2000; (14,000–4,000 cm1) region is often called the
van Dishoeck 2004). In brief, ISO showed con- near-infrared, the 2.5–25 mm (4,000–400 cm1)
clusively that space is full of the building blocks region is called the mid-infrared, and the
of life. 25–1,000 mm (400–10 cm1) region is called
the far-infrared.

Overview
Cernicharo J, Crovisier J (2005) Water in space: the water
world of ISO. Space Sci Rev 119:29–69 Many organic and inorganic materials absorb
Cesarsky C, Salama A (2005) ISO science legacy. certain frequencies of infrared light. The absorp-
Springer, Dordrecht
tion is caused by vibrations or rotations of mole-
Cox P, Kessler MF (1999) The universe as seen by IS-
O. ESA SP-427. ESTEC, Noordwijk, 1090 pp cules, at frequencies corresponding to the
Ehrenfreund P, Charnley SB (2000) Organic molecules in infrared region of the electromagnetic spectrum.
the interstellar medium, comets and meteorites: a voy- Therefore, information about molecular species
age from dark clouds to the early Earth. Annu Rev
and structure can be obtained by irradiating sam-
Encrenaz T (2008) Water in the solar system. Annu Rev ples in the infrared frequency range and subse-
Astron Astrophys 46:57–87 quently observing the absorption spectra or by
First ISO Results (1996) A&A 315:L27–L400 observing the corresponding emission spectra if
Kessler MF et al (1996) The Infrared Space
Observatory (ISO) mission. Astron Astrophys 315:
the intensities exceed that of the background
L27–L31 radiation. Vibrational transitions in the infrared
Van Dishoeck EF (2004) ISO spectroscopy of gas for some typical functional groups occur in the
and dust: from molecular clouds to regions of 3,600–2,500 cm1 (OH stretching),
protoplanetary disks. Annu Rev Astron Astrophys
3,100–2,800 cm1 (CH stretching),
42:119–167
1,800–1,650 cm1 (C=O stretching), and
1,600 cm1 (aromatic C=C stretching)
(Socrates 2001).
In the laboratory the spectral intensity is
Infrared Spectroscopy usually reported as a transmittance (T) or absor-
bance (A). The transmittance is defined as the
Yoko Kebukawa ratio of infrared energy passing through a
Department of Natural History Sciences, sample (I) and irradiated infrared energy
Hokkaido University, Sapporo, Japan (background; I0):
T ¼ I=I 0 ;
Keywords
which is usually shown in % transmittance (%T).
Fourier transform infrared microspectroscopy; The absorbance is defined as
Fourier transform infrared spectroscopy; FTIR;
A ¼ log10 T ¼ log10 ðI=I 0 Þ:
Infrared microscopy
Initial Mass Function 1197

The absorbance is linearly correlated with the
concentration of the component of interest, a Bellamy LJ (1975) The infrared spectra of complex mol-
relation known as the Lambert-Beer law: ecules, 3rd edn. Chapman and Hall, London
Colthup NB, Daly LH, Wiberley SE (1990) Introduction
A ¼ edC; to infrared and Raman spectroscopy, 3rd edn. Aca-
demic, San Diego
Griffiths PR, de Haseth JA (1986) Fourier transform infra-
where e is the molar absorptivity, d is the thick- red spectrometry. Wiley, New York
ness of the sample, and C is the concentration of Holman HYN, Martin MC (2006) Synchrotron radiation
the component of interest. The infrared absorp- infrared spectromicroscopy: a non-invasive chemical
probe for monitoring biogeochemical processes. Adv
tion intensity (or integral absorption intensity), Agron 90:79–127
with appropriate baseline correction, can be Silverstein RM, Webster FX, Kiemle D (2005) Spectro-
therefore used for quantitative analyses. metric identification of organic compounds, 7th edn.
Today, Fourier transform infrared spectros- Wiley, New York
Socrates G (2001) Infrared and Raman characteristic
copy (FTIR) is more commonly used than tradi- group frequencies, 3rd edn. Wiley, Chichester
tional monochromatic infrared spectroscopy. The
instrumentation of FTIR usually consists of a
light source, an interferometer (instead of a I
monochromator), a detector, and a computer sys-
tem. In FTIR, the time domain spectrum Initial Mass Function
(interferogram) obtained by infrared light passing
thorough an interferometer is converted to the Leticia Carigi
frequency-domain spectrum by a Fourier trans- Instituto de Astronomı́a, Universidad Nacional
form. A microscope is often combined with an Autónoma de México, México, DF, Mexico
FTIR for micro-to-millimeter-sized samples and
is useful for spatial characterization. Most
recently, a synchrotron radiation light source Keywords
has been used instead of a conventional ceramic
light source. The greater brightness of a synchro- Evolution of galaxies; Low mass stars; Massive
tron source enables a better spatial resolution stars; Solar neighborhood; Star formation
(close to the expected diffraction limit according
to the wavelength) and a higher signal-to-noise
ratio than ceramic light sources (Holman and Acronyms
Martin 2006).
Infrared spectroscopy can be applied to solid, IMF
liquid, and gas samples. Infrared spectroscopy is
used in astronomical observations of the solar
system and the ▶ interstellar medium, e.g., for Definition
identifying constituents of planetary atmospheres
and surfaces, and the constituents of ▶ molecular The initial mass function is the relative number of
clouds in the Milky Way and external galaxies. stars, as a function of their individual initial mass,
that forms during a single star-forming episode.
See Also
Overview
▶ Infrared Astronomical Satellite
▶ Infrared Astronomy The initial mass function (IMF) is usually expressed
▶ Infrared Space Observatory as a power law, IMF = constant ma,
1198 Initial Mass Function, Origin of
where m is the initial star mass and a is the slope of See Also
the logarithmic plot. The IMF is defined over a
large interval of masses, from the most massive ▶ Solar Neighborhood
stars to the lowest mass stars, created in a single ▶ Star Formation, Theory
▶ star formation burst. ▶ Stellar Evolution
The majority of the empirical IMFs has
been inferred from star counts observed in the
solar neighborhood. This region has had a
long history of star formation bursts, including Briceño C et al (2002) The initial mass function in the
stars of different ages. Hence, to determine taurus star-forming region. Astrophys J 580:317–335
the true IMF, it is necessary to correct the Carigi L, Hernandez X (2008) Chemical consequences of
actual count, assuming how the star low star formation rates: stochastically sampling the
initial mass function. MNRAS 390:582–594
formation rate has changed over time, stellar Kroupa P (2001) On the variation of the initial mass
ages, the number of binary or multiple function. MNRAS 322:231–246
stellar systems, the Galactic age, and some other Kroupa P, Tout C, Gilmore G (1993) The distribution of
factors. low-mass stars in the Galactic disc. MNRAS
262:545–587
The first and still most popular IMF is that Salpeter EE (1955) The luminosity function and stellar
obtained by Salpeter (1955), with slope evolution. Astrophys J 121:161–167
a = 2.35, over the entire mass interval. It is
important to mention that Salpeter’s original
calculation was for stars between 0.4 and 10 M
and the astronomical community has often
extrapolated it without careful consideration. Initial Mass Function, Origin of
When Salpeter’s IMF is extrapolated to m < 0.4
M, the number of very low mass stars is Gilles Chabrier
one or two orders of magnitude larger than Centre de Recherche Astrophysiue de Lyon,
the observed value. In the literature, there are Ecole Normale Supérieure de Lyon, Lyon,
many different IMFs (e.g., Kroupa et al. 1993), France
all of them with several slope values; that is,
they suggest different slopes for different mass
intervals. The disagreement is larger for the low Keyword
mass stars, due to their intrinsically low
luminosity. Star formation
It is known that Nature tends to form
much more low mass stars than high mass stars,
and this can be seen for the values proposed for a Overview
(larger than +2). For instance, Kroupa
et al. (1993) predict approximately 4,000 stars It was first recognized by Richard Larson in 1981
of 1 M and 40 stars of 10 M for each star of that ▶ molecular clouds are dominated at large
100 M. scales by strong turbulent motions. ▶ Turbulence
Since stellar properties mainly depend on their thus determines the structure of all density
initial masses, the IMF is fundamental to the regimes in the interstellar gas and in star-forming
chemical and spectral evolution of star clusters molecular clouds (MacLow and Klessen 2004).
and galaxies. The sources of this turbulence are multiple and
It is very important to note that the universality can be due to convergent flows in the interstellar
of IMF remains as an unsolved problem and so medium, supernova explosions, galactic shear, or
does the behavior of the IMF for substellar even more simply to star formation itself. Given
masses (m < 0.08 M). the low density of the medium, the turbulent flow
Initial Mass Function, Origin of 1199
is highly compressible and cascades from large to emergence of a lognormal form for the PDF, at least
small scales by shock waves. Typical Mach num- for the aforementioned typical Mach values, both in
bers of this turbulence in star-forming molecular supersonic hydrodynamic (e.g., Padoan and
clouds range from about 3 to 8 under usual Milky Nordlund 2002; Federrath et al. 2010; Schmidt
Way-like conditions, although more extreme et al. 2010) and MHD turbulence (Lemaster and
values can be found in starburst environments. Stone 2008; Price et al. 2011). Although departure
Observations show a power law dependence from such a lognormal form does occur under some
between the typical turbulent velocity dispersion, circumstances, due to the intermittency of turbu-
s, in molecular clouds and the size of the lence or the nonisothermality of the gas, it remains
cloud, L, a relation known as one of the Larson’s a reasonable approximation for the aforementioned
relations, s/LZ (see Hennebelle and Falgarone typical Mach values.
2013 for a recent review). The power index of this Interestingly enough, observational determi-
relation, Z, is in fact tightly connected to the nations of the stellar initial mass function, defined
index of the turbulence velocity power spectrum, originally as the number of stars per logarithmic
n, with Z = (n3)/2 (e.g., Hennebelle and Cha- mass interval, dN/dlogM, in various Galactic
brier 2008). Numerical simulations of compress- environments, in the field, in young clusters,
ible turbulence show that this index is of the order and in star-forming regions, confirm that the
I
of n ~ 3.8, in three dimensions (Beresnyak IMF exhibits a power law tail for masses above
et al. 2005; Kritsuk et al. 2007), typically between about a solar mass, the well-known Salpeter IMF
the Kolmogorov value n = 5/3 for incompress- (Salpeter 1955), but rolls over below about 0.8
ible turbulence and the Burgers value n = 4 for solar masses. The IMF is globally well described
shock-dominated turbulence. As discussed by a combination of this power law tail and a
below, this index plays a key role in the determi- lognormal form, extending well into the brown
nation of the slope of the ▶ initial mass function dwarf regime (Chabrier 2003, 2005). Most inter-
(IMF). estingly, the observations of gravitationally
bound prestellar cores in various star-forming
regions, as revealed in particular by the Herschel
Key Research Findings satellite (André et al. 2010), called the core mass
function (CMF), precursor of the stellar IMF,
The statistics of the gas density fluctuations created displays as well a Salpeter power law tail turning
by turbulence is captured by the so-called probabil- down to a lognormal shape at low masses, closely
ity density function (PDF). A key property of tur- resembling the Chabrier IMF. All these numeri-
bulence is that in good approximation, the PDF of cal experiments and observations thus strongly
the logarithm of the density (log r) is Gaussian, suggest that star formation and turbulence are
implying that the PDF is lognormal. Such a distri- closely linked and that the IMF is imprinted by
bution is expected from the application of the cen- turbulence in the cloud at the very early stages of
tral limit theorem to a hierarchical density field the star formation process.
generated by multiplicative processes, such as
shocks (Vázquez-Semadeni 1994). Such a lognor- Theory of the Stellar Initial Mass Function
mal behavior for density fluctuations has received Several attempts have been made to develop a
both observational and numerical support. Obser- theory linking turbulence and the origin of the
vationally, the 3D lognormal shape of the PDF is IMF. The first attempt to demonstrate the link
reflected by the projected (2D) power spectrum between the lognormal form of the turbulence
column density of molecular clouds, measured PDF and the distribution of Jeans unstable
from dust extinction maps (Kainulainen prestellar cores was made by Padoan and
et al. 2009; Lombardi et al. 2010; Brunt Nordlund (2002). Although indeed illustrating
et al. 2010). On the other hand, various numerical the relationship between these two quantities,
simulations of compressible turbulence do show the this theory, however, is based on several ad hoc
1200 Initial Mass Function, Origin of
assumptions and cannot properly reproduce the Jeans mass will gravity take over, yielding the
Salpeter slope of the IMF in the pure hydrody- collapse of the corresponding mass reservoirs,
namic case (without magnetic field), in contrast to providing the initial seeds for the massive gravita-
what is obtained in numerical simulations. Later tionally bound prestellar cores.
on, Hennebelle and Chabrier (2008, 2009), using In this theory, this often called “turbulent sup-
the analogy between cosmological and turbulence port” should thus not be confused with a static
Gaussian density fluctuations, have extended the support, equivalent to a pressure, but should
formalism of Press and Schechter (1974), origi- rather be seen as a random process selecting at
nally developed to determine the mass function of the very early stages of the star formation process
collapsing dark halos in the primordial universe, to the pieces of gas that will be dominated by grav-
the star formation process. In that case, however, ity and will thus collapse, from the ones that will
the random field of density fluctuations is no lon- be dispersed. Eventually, turbulence will of
ger uniform, as for the primordial universe, but course quickly dissipate, and by the time they
lognormal, as mentioned above. Combining this are observed, the bound cores will exhibit only
distribution of initial density fluctuations with the thermal inner motions, but turbulence will have
virial condition for gravitational collapse, which been the initial triggering mechanism. This key
states that only fluctuations within which gravity role of turbulent motions, characterized by the
dominates over all other sources of support are velocity power spectrum index mentioned previ-
gravitationally unstable, this theory reproduces ously, is mandatory to obtain the proper Salpeter
all of the key features of the observed CMF and, slope at high masses for the IMF, the slope being
by extrapolation, of the stellar IMF, including the directly related to this index (Hennebelle and
mass distribution of brown dwarfs, for appropriate Chabrier 2008; Chabrier and Hennebelle 2010).
density and Mach number conditions. In this the- This concept of turbulent Jeans mass for collapse
ory, the impact of turbulence is twofold. First of has received some support from numerical simu-
all, turbulence-induced shocks lead to a field of lations aimed at exploring this issue (Schmidt
overdensities, and fluctuations of the underlying et al. 2010). Later on, also using analytical tools
velocity field follow a Kolmogorov-like scale originally developed for cosmology, Hopkins
dependence. Fluctuations dense enough to (2012) generalized the concept of Hennebelle
become gravitationally unstable provide the and Chabrier to larger, galactic scales, which
seeds for the future collapsing prestellar cores, yields the mass function of giant molecular
including pre-brown dwarf cores at very small clouds in galactic disks. As in the Hennebelle-
scales. The more powerful the turbulence, i.e., Chabrier theory, the stellar IMF in the Hopkins
the higher the large-scale typical Mach number, formalism is entirely determined by one single
the larger the number of small-scale overdensities, parameter, namely, the turbulent velocity power
thus, of low-mass cores. On the other hand, turbu- spectrum index.
lence constantly sweeps away gas reservoirs
which, otherwise, would become Jeans unstable
and collapse. As a consequence, mass reservoirs Future Directions
can grow much larger than in the absence of tur-
bulent motions, until eventually they exceed a The striking resemblance between the prestellar
mass defined as the turbulent, as opposed to purely core mass function and the final stellar IMF,
thermal, Jeans mass. In other words, overdense added to the growing evidence for this latter
regions with lower masses than the turbulent being the combination of a Salpeter power law
Jeans mass will be dispersed back into the flow at large masses and a lognormal form below
under the action of turbulence within a turbulent about one solar mass, clearly points to two fun-
crossing time, before they become dominated by damental results for the origin of the IMF. First of
gravity. Only in regions exceeding the turbulent all, this latter seems to be imprinted in the cloud
Initial Mass Function, Origin of 1201
at the very early stages of star formation and be References and Further Reading
determined by the very conditions in the cloud,
density, temperature, and large-scale Mach num- André P et al (2010) From filamentary clouds to prestellar
cores to the stellar IMF: initial highlights from the
ber. Second, observations and numerical simula-
Herschel Gould Belt Survey. Astron Astrophys
tions show that compressible turbulence, with or 518:102–106
without the presence of a magnetic field, leads to Beresnyak A, Lazarian A, Cho J (2005) Density scaling
a roughly lognormal distribution of overdensities and anisotropy in supersonic magnetohydrodynamic
turbulence. Astrophys J 624:93–96
mirroring the shape of the IMF. As discussed
Brunt C, Federrath C, Price D (2010) A method for
above, this concept of turbulence-induced frag- reconstructing the PDF of a 3D turbulent density field
mentation for the origin of the IMF can now rely from 2D observations. MNRAS 405:56–60
on a sound theoretical support, which shows that Chabrier G (2003) Galactic stellar and substellar initial
mass function. Publ Astron Soc Pac 809:763–795
the very shape of the IMF is entirely determined
Chabrier G (2005) The initial mass function: from
by the density and velocity power spectra of Salpeter 1955 to 2005. Astrophys Space Sci Libr
compressible turbulence, which are universal 327:41
properties. Quantitatively speaking, however, Chabrier G, Hennebelle P (2005) Dimensional argument
for the impact of turbulent support on the stellar initial
the IMF, notably its extension in the low-mass
mass function. Astron Astrophys 534:106–110
domain, depends on the essential properties of the Federrath C, Roman-Duval J, Klessen R, Schmidt W, Mac I
cloud, density, temperature, and large-scale Low M-M (2010) Comparing the statistics of interstel-
Mach number. Under standard Milky Way-like lar turbulence in simulations and observations. Sole-
noidal versus compressive turbulence forcing. Astron
molecular cloud conditions, these properties
obey Larson’s relations and display rather similar Hennebelle P, Chabrier G (2008) Analytical theory for the
values, leading to a rather universal IMF. It is not initial mass function: II. Properties of the flow.
excluded and is indeed predicted by the above Astrophys J 684:395–410
Hennebelle P, Chabrier G (2009) Analytical theory for the
theories, however, that under quite different con-
initial mass function: CO clumps and prestellar cores.
ditions, for instance, in extreme environments, Astrophys J 702:1428–1442
the IMF might differ quantitatively from the one Hopkins P (2012) The stellar initial mass function, core
inferred in the galactic neighborhood, in particu- mass function and the last-crossing distribution.
MNRAS 423:2037–2044
lar for the peak position and the extension in the
Kainulainen J, Lada C, Rathborne J, Alves J (2009) The
low-mass regime. If the theory is correct, how- fidelity of the core mass functions derived from dust
ever, the IMF should qualitatively remain the column density data. Astron Astrophys 534:106–110
same, with a high-mass power law tail rolling Kritsuk A, Norman M, Padoan P, Wagner R (2007) The
statistics of supersonic isothermal turbulence.
down to a lognormal form below about the char-
acteristic thermal Jeans mass of the medium. Larson R (1981) Turbulence and star formation in molec-
Future observations, both in the galactic ular clouds. MNRAS 194:809–826
neighborhood and in extragalactic or widely dif- Lemaster M, Stone J (2008) Density probability distribu-
tion functions in supersonic hydrodynamic and MHD
ferent environments, should allow us to test fur-
turbulence. Astrophys J 682:97–100
ther these theories and verify the robustness of Lombardi M, Lada C, Alves J (2010) 2MASS wide
this general paradigm for the origin of the stellar field extinction maps. III. The Taurus, Perseus, and
initial mass function. California cloud complexes. Astron Astrophys
512:67–81
MacLow M-M, Klessen R (2004) Control of star forma-
tion by supersonic turbulence. Rev Mod Phys
See Also 76:125–194
Padoan P, Nordlund A (2002) The stellar initial mass
▶ Fragmentation of Interstellar Clouds function from turbulent fragmentation. Astrophys
J 576:870–879
▶ Initial Mass Function Press W, Schechter P (1974) Formation of galaxies and
▶ Star Formation, Observations clusters of galaxies by self-similar gravitational con-
▶ Star Formation, Theory densation. Astrophys J 187:425–438
1202 Insoluble Organic Matter
Price D, Federrath C, Brunt C (2011) The density materials. Even though various organic com-
variance-mach number relation in supersonic, isother- pounds including amino acids, carboxylic acids,
mal turbulence. Astrophys J 727:21–26
Salpeter E (1955) The luminosity function and stellar and polycyclic aromatic hydrocarbons (PAHs)
evolution. Astrophys J 121:161–167 have been identified in meteorites, particularly
Schmidt W, Kern S, Federrath C, Klessen R (2010) Numer- in carbonaceous chondrites, these solvent-
ical and semi-analytic core mass distributions in super- extractable organic compounds are only
sonic isothermal turbulence. Astron Astrophys 516:25–39
Vázquez-Semadeni E (1994) Hierarchical structure in 10–30 % of the total organic carbon. Most mete-
nearly pressureless flows as a consequence of self- oritic organic matter is insoluble in water, acids,
similar statistics. Astrophys J 7(423):681–692 or organic solvents. This insoluble organic matter
is used to be called kerogen-like material
(cf. kerogen is a solvent-insoluble organic matter
in terrestrial rocks) or macromolecular organic
Insoluble Organic Matter matter. Now, it is generally called IOM.
For the ▶ Murchison meteorite, which is rela-
Hiroshi Naraoka tively enriched in soluble organic compounds,
Department of Earth and Planetary Sciences, the IOM consists of more than 90 % of the mete-
Kyushu University, Fukuoka, Japan oritic organic carbon with minor amounts of
hydrogen, nitrogen, oxygen, and sulfur. Although
the chemical structure of IOM has not been
Keywords defined accurately yet, it has aromatic hydrocar-
bon cores (two to four aromatic rings) with
Alteration; Extraterrestrial; Insoluble; Organic short-chain aliphatic hydrocarbons. IOM from
matter; Unusual isotopic composition various meteorites has a wide variation in
chemical composition. The H/C ratio of chon-
dritic IOM varies from <0.1 to 0.7, the more
Synonyms metamorphosed meteorite having a lower H/C
ratio. Therefore, the H/C ratio is a sensitive
IOM; Kerogen-like matter indicator for thermal processing during the
meteorites’ history. 13C-NMR and ▶ XANES
studies have also revealed various chemical
Definition bonding and functional groups, including aro-
matic and aliphatic structures with carbonyl, car-
Insoluble organic matter (IOM) is a major compo- boxyl, and aldehyde as well as amide and nitrile
nent of organic matter found in primitive extrater- groups. As the Murchison IOM generates abun-
restrial materials such as carbonaceous meteorites. dant PAHs and acetic acid (ca. four times higher
Most of this is organic carbon with lesser amounts compared to solvent-extractable acetic acid) dur-
of hydrogen, nitrogen, oxygen, and sulfur forming ing hydrous pyrolysis, the IOM may be an
aromatic hydrocarbon cores (two to four aromatic important precursor for organic compounds in
rings) and short-chain aliphatic hydrocarbons with meteorites.
various functional groups including nitrile, car- The most primitive IOM also possesses
boxyl, and ether. Unusual isotopic signatures such unusual isotopic signatures such as extreme d-
as extreme d- and 15N-enrichment are observed. and 15N-enrichment, which have not been
observed in solar system processes. Such isotopic
behavior suggests that meteoritic organic matter
Overview may have originated from interstellar processes.
However, the unusual isotope signals are dimin-
Insoluble organic matter (IOM) is a major con- ished or lost during the thermal and aqueous
stituent of organic matter in extraterrestrial alteration on meteorite parent bodies.
Intelligence 1203
See Also
Institut for Rumforskning og -
▶ Carbonaceous Chondrite teknologi på Danmarks Tekniske
▶ Carbonaceous Chondrites, Organic Chemistry of Universitet
▶ Murchison ▶ DTU Space, Denmark
▶ XANES

Intellect
Alexander CMO’D, Fogel M, Yabuta H, Cody GD
(2007) The origin and evolution of chondrites recorded ▶ Intelligence, Evolution of
in the elemental and isotopic compositions of their ▶ Intelligence
macromolecular organic matter. Geochim Cosmochim
Acta 71:4380–4403
Alexander CMO’D, Newsome SD, Fogel ML, Nittler LR,
Busemann H, Cody GD (2010) Deuterium enrich-
ments in chondritic macromolecular
material – implications for the origin and evolution
Intelligence I
of organics, water and asteroids. Geochim Cosmochim
Acta 74:4417–4437 Lori Marino
Busemann H, Young AF, Alexander CMO’D, Hoppe P, Emory Centre for Ethics, Emory University,
Mukhopadhyay S, Nittler LR (2006) Interstellar chem-
istry recorded in organic matter from primitive mete-
Atlanta, GA, USA
orites. Science 312:727–730
Cody GD, Alexander CMO’D (2005) NMR
studies of chemical structural variation of insoluble Keywords
organic matter from different carbonaceous
chondrites groups. Geochim Cosmochim Acta
69:1085–1097 Cognition; Complexity; Brain; fi;
Huang Y, Alexandre MR, Wang Y (2007) Encephalization; Neuroanatomy
Structure and isotopic ratios of aliphatic side chains
in the insoluble organic matter of the Murchison car-
bonaceous chondrites. Earth Planet Sci Lett
259:517–525 Synonyms
Kerridge JF, Chang S, Shipp R (1987) Isotopic character-
ization of kerogen-like material in the Murchison car- Cleverness; Cognition; Intellect
bonaceous chondrite. Geochim Cosmochim Acta
51:2527–2540
Oba Y, Naraoka H (2006) Carbon isotopic composition of
acetic acid generated by hydrous pyrolysis of macro- Definition
molecular organic matter from the Murchison meteor-
ite. Meteorit Planet Sci 41:1175–1181
Oba Y, Naraoka H (2009) Elemental and isotopic behavior
Intelligence is a term referring to a range of
of macromolecular organic matter from CM chon- abilities that have to do with how an individual
drites during hydrous pyrolysis. Meteorit Planet Sci (or collective entity) acquires, processes, stores,
44:943–954 analyzes, and acts upon information and situa-
Remusat L, Derenne S, Robert F, Knicker H (2005) New
pyrolytic and spectroscopic data on Orgueil and Mur-
tions. It includes capacities such as memory,
chison insoluble organic matter. A different origin learning, problem solving, abstract thinking, cre-
than soluble? Geochim Cosmochim Acta ativity, and behavioral flexibility. Intelligence is,
69:3919–3932 by nature, a fuzzy concept. That is, there are no
Remusat L, Palhol F, Robert F, Derenne S, France-Lanord
C (2006) Enrichment of deuterium in insoluble organic
strict boundaries on it and there is no scientific
matter from primitive meteorite: a solar system origin? consensus on its definition. Intelligence is often
Earth Planet Sci Lett 243:15–25 referred to as cognitive complexity.
1204 Intelligence
Overview mechanisms of learning and memory are shared

across all species, including humans. There are
In the domain of ▶ SETI, intelligence has been fundamental capacities which even simple ner-
operationalized as the presence of a technology vous systems allow which are ubiquitous among
detectable from Earth. In the broader framework all motile life forms on earth. And there is a deep
of astrobiology, however, there is no need to limit continuity in the domain of intelligence across
the study of intelligence to a strict technical cri- species. Likewise, in metazoans, intelligence is
terion. The study of intelligence in the often correlated with other psychological capac-
astrobiological paradigm involves applying our ities such as self-awareness, emotional complex-
knowledge and understanding of the myriad ity, and personality structure. Even “high-level”
intelligences on earth to develop scientific cognitive abilities once thought to be exclusively
hypotheses about extraterrestrial intelligence. possessed by humans, such as for planning, basic
Intelligence is a reflection of the kinds of cogni- mathematics, self-recognition, and a technologi-
tive adaptations each species has had to make in cal culture, are not unique to our species.
its evolutionary history and is, thus,
multidimensional, interacting with many other
adaptive features, such as sensory-perceptual See Also
and motor abilities.
Intelligence across species on this planet is ▶ Astrobiology
measured in a variety of ways. These include ▶ Bioastronomy
experimental studies of learning, memory, and ▶ Biological Evolution
other aspects of cognition in animals. It also ▶ Complexity
involves determining what cognitive capabilities ▶ Complex Organisms
are evinced in animals in a natural setting. There ▶ Darwinian Approaches
are trade-offs with the validity of both ▶ Drake Equation
approaches. Several neuroanatomical measures ▶ Ecological Niche
are used as proxies for intelligence across spe- ▶ Exobiology
cies, including relative brain size (also known as ▶ Intelligence, Evolution of
encephalization quotient), the volume of specific ▶ Search for Extraterrestrial Intelligence
structures in the brain, and nerve cell density and ▶ SETI
connectivity. Some combination of all of the ▶ Species
above features within the categories of brain
size and organization forms the neurological
basis of the phenomenon we know as intelli-
gence. Correlations among various measures of References and Further Reading
brain size, cognitive capacity, problem solving,
and behavioral ecology, e.g., social complexity, Allman J (1999) Evolving brains. Scientific American
Library, New York
dietary complexity, have been found in many Marino L (2004) Objectivity in the study of intelligence:
groups of animals across phyla. These patterns the cornerstone of new methods and discoveries. In:
of association point to possible scenarios about Norris R, Oliver C, Stootman F (eds) Proceedings of
the causal relations between intelligence and the bioastronomy 2002 symposium: life among the stars.
Astronomical Society of the Pacific, San Francisco,
physical and social environment. The challenge pp 163–193
is to determine how all of these features relate to Marino L (2011) Cetaceans and primates: convergence in
each other to form the range of intelligence that intelligence and self-awareness. J Cosmol
exists both within and across species on earth. 14:1063–1079
Roth G, Dicke U (2005) Evolution of the brain and intel-
Basic cognitive processes forming the funda- ligence. Trends Cogn Sci 9:250–257
mental groundwork for intelligence in animals Striedter GF (2005) Principles of brain evolution. Sinauer
are found in unicellular organisms as Associates, Sunderland
Intelligence, Evolution of 1205
branch lengths, often, are used to represent time.

Intelligence, Evolution of Phylogenetic comparative methods are then
employed, using the tree, to test for patterns of
Lori Marino evolutionary change from which we can draw
Emory Centre for Ethics, Emory University, inferences about the evolutionary process.
Atlanta, GA, USA The kinds of data used to develop a phyloge-
netic tree range from phenotypic, behavioral, and
anatomical traits to genotypic characteristics.
Keywords Comparative phylogenetic analysis provides a
way to, for instance, infer the ancestral state of
Cognition; Complexity; Brain; Central nervous a given trait, e.g., toolmaking or mirror
system; Encephalization; fi self-recognition, or whether there is evidence
for convergent evolution in some component of
intelligence. The actual data compared against a
Synonyms phylogenetic framework derive from a variety of
approaches.
Cleverness; Cognition; Intellect Research on intelligence in living animals is
I
typically conducted using two approaches:
laboratory-based experimental studies and obser-
Definition vations in a naturalistic ethological setting. There
are trade-offs with the validity of both
Intelligence, those features of an organism that approaches, and this is why both must be com-
mediate the way in which that organism repre- bined to understand intelligence in a comparative
sents, processes, and acts upon information, is a evolutionary framework. The strength of
biological trait subject to the same evolutionary laboratory-based experimental tests of cognition
principles that shape all other biological features is that they can be well controlled for extraneous
of organisms on this planet. The evolution of variables, providing a way to test specific hypoth-
intelligence refers to the pattern, distribution, eses about the existence of certain capacities in a
and mechanisms of neurobiological, cognitive, species. The weakness in this approach is in the
and behavioral characteristics within and across questionable generalizability to a natural setting.
species over time. The question of the evolution The challenge for laboratory-based approaches
of intelligence is often framed as one of cognitive when studying animal intelligence is to devise
(psychological) complexity. tasks which are species appropriate, i.e., tap into
the natural proclivities and sensitivities of the
species. Yet, despite the artificial setting of a
Basic Methodology laboratory, many members of other species
score high marks on human-devised tests of intel-
The main task in studying the evolution of intelligence and even exceed human performance in
ligence is to find ways to reconstruct the past and specific domains. For instance, chimpanzees far
place the cognitive characteristics of modern spe- exceed human performance on numerical short-
cies in the proper temporal and phylogenetic con- term memory tests under equivalent laboratory
text. This begins with a well-supported conditions, i.e., using a touch-sensitive computer
phylogenetic tree, i.e., a branching diagram of screen (Inoue and Matsuzawa 2007).
species showing their inferred evolutionary rela- The other approach, known as cognitive ethol-
tionships based on physical and genetic traits. ogy, assesses abilities of animals in their natural
Connected taxa on the tree are implied to have habitat and therefore frames intelligence more
descended from a common ancestor. Nod

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Are we alone? Long an object of speculation or fiction, if not heresy, this

chance provided enough opportunities for selection to be optimizing and,

14 January 2011 Christian de Duve

presently growing exponentially, with no end in sight. Encyclopedias now

Simpson famously noted shortly thereafter that Exobiology was a science

where N is the number of civilizations in our galaxy for which communica-

The Encyclopedia of Astrobiology serves as the key to a common under-

A brief note is warranted about how we constructed the Encyclopedia.

Muriel Gargaud CNRS-Universite de Bordeaux, Laboratoire

William M. Irvine University of Massachusetts, Amherst, MA, USA

Ricardo Amils Departamento de Biologia Molecular, Universidad

Alessandro Airo Institut f€ur Geologische Wissenschaften Tektonik und

Francis Albarède Ecole Normale Supérieure de Lyon, Lyon, France

Yann Alibert Space Research and Planetary Sciences, Physics Institute,

Philippe André Laboratoire AIM, IRFU/Service d’Astrophysique, CEA

Kristin Bartik Université libre de Bruxelles, Brussels, Belgium

Hugues Bersini IRIDIA, Université Libre de Bruxelles, Brussels, Belgium

Carlos Briones Centro de Astrobiologı́a (CSIC/INTA), Consejo Superior de

Therese Encrenaz LESIA, Observatoire de Paris - Section de Meudon,

Fernando B. Figueiredo CITEUC, University of Coimbra, Coimbra,

Felipe Gomez Centro de Astrobiologı́a (CSIC/INTA), Instituto Nacional de

José Pascual Abad Facultad de Ciencias, Departamento de Biologı́a Molec-

Luc André Department of Earth Sciences, Royal Museum of Central Africa,

Sylvain Bernard Laboratoire de Minéralogie et de Cosmochimie du

Hugues Bersini IRIDIA, Université Libre de Bruxelles, Brussels, Belgium

Bruno Bézard LESIA, Observatoire de Paris, Meudon, France

Jean-Pierre Bibring Institut d’Astrophysique Spatiale, Université Paris

John H. Black Department of Earth and Space Sciences, Chalmers Univer-

Donna Blackmond The Scripps Research Institute, La Jolla, CA, USA

Jeffrey Blanchard Biology Department, University of Massachusetts

Laurent Boiteau Institut des Biomolécules Max Mousseron, UMR5247

Tyler Bourke Square Kilometre Array Organisation, Macclesfield,

Jessica C. Bowman School of Chemistry and Biochemistry, Georgia Insti-

Samuel A. Bowring Department of Earth, Atmospheric and Planetary

Maud Boyet Université Blaise Pascal, Clermont-Ferrand, France

André Brack Centre de Biophysique Moléculaire CNRS, Orléans Cedex 2,

Robert Braun Square Kilometre Array Organisation, Macclesfield,

Doris Breuer German Aerospace Center (DLR), Institute of Planetary

Carlos Briones Centro de Astrobiologı́a (CSIC/INTA), Consejo Superior de

Gilles Bruylants Engineering of Molecular NanoSystems, Universté Libre

Sergey Bulat NRC ‘Kurchatov institute’, Petersburg Nuclear Physics Insti-

Vincent Busigny Institut de Physique du Globe de Paris, Paris, France

Gary R. Byerly Department of Geology & Geophysics, Louisiana State

Michel Cabane LATMOS/IPSL B102/T45-46, Université Pierre et Marie

Gregory Chambon Faculty of Arts and Humanities, University of Nantes,

Bradley De Gregorio Materials Science and Technology Division, U.S.

Sylvia Ekström Observatoire Astronomique de l’Université de Genève,

José Carlos Gaspar Institute of Geosciences, University of Brası́lia,

udel Department of Astrophysics, University of Vienna, Vienna,

Pentti Hölttä Department of Geosciences and Geography, University of

Anders Johansen Lund University, Lund, Sweden

Kurt O. Konhauser Department of Earth and Atmospheric Sciences,

Akira Kouchi Institute of Low Temperature Science, Hokkaido University,

Ramanarayanan Krishnamurthy Chemistry, The Scripps Research